Reprinted from ANNALS ENTOMOLOGICAL SocJETY OF AMERICA Vol. 49 No. 1. January, 1956 THE GENUS IN NORTH AMERICA (EPHEMEROPTERA: ) 1

LEWIS BERNER Department of Biology, University of Florida, Gainesville, Fla.

With the discovery of a new species of N eo­ same paper she mentioned that another species, ephemera, it seems advisable to review our represented only by nymphs, was at hand, and present knowledge of all of the species of the this same nymph was again reported by her in genus and to construct keys for their separation. 1935 as occurring in Florida. In 1935, the It is certainly likely that as our knowledge of the genus was transferred to the family , of North America is enlarged additional subfamily Neoephemerinae (Needham, Traver, species of N eoephemero will be described; such a and Hsu, 1935). The second species of Oreionthus summary may, therefore, serve as a guide for was described from Florida nymphs by Traver future work. (1937) but was not given a specific name, simply McDunnough (1925) described Neoephemero being designated as Oreionthu; sp. No. I. from eight males that were collected by G. S. Oreianthus was next discussed in "The May­ Walley at Laprairie, Quebec. In his paper, flies of Florida" (Berner, 1950) where it was McDunnough gave no information relative to said that phylogenetically the genus probably the phylogenetic relationship of the genus nor represented a distinct family. did he discuss conditions under which the speci­ Burks (1953) established the family Neo­ mens were taken. Ulmer (1933) was the first to ephemeridae to include the genera N eoephemera give N eoephemero a family placement when he and N eoephemeropsis and at the same time included it as one of the genera of the Ephemeri­ synonymized Oreianthus with N eoephemera. When dae. Subsequently, Needham, Traver, and Hsu Ulmer (1939) described N eoephemeropsis from (1935) erected a new subfamily, Neoephemerinae, Sumatra and Java, he placed it in the family to include McDunnough's genus as well as Potamanthidae and stated that it was close to Oreionthus, a genus described by Traver in 1931. N eoephemera; however, Burks claims that N eo­ In this latter report, it was stated that the ephemeropsis almost certainly belongs in the nymph of N eoephemero was unknown and that family Neoephemeridae, a conclusion with which the distribution of the genus was Nearctic; I am in agreement. Oreionthus was said to be Holarctic, and in the With the description of N eoephemero youngi Nearctic to occur in the southeastern United Berner (1953), the status of 1 •aver's Oreionthus States. sp. No. I was clarified. Subsequently, I have The description of Oreionthus was based on discovered a fourth species in the genus, the specimens collected and reared in 1929 and 1930. description of which is given below. Traver (1931), in her description, indicated that As presently understood, the genus in North the wing venation of the adult showed America is distributed from southeastern Canada that it should be placed in the subfamily Ephem­ southward through the Appalachians to Florida erinae even though the nymph, other than in and westward is known from Michigan. Nymphs size, resembles that of the genus Caenis. In and adults are rarely found in collections and discussing the relations of Oreianthus to other records of the occurrence of the genus are sparse. forms, Traver pointed out that, based on a The known distribution of each species will be study of illustrations, a nymph from the Garonne discussed separately. River in France, which was described by Joly I am indebted to Dr. Henry Dietrich, Cornell (1870) as Caenis maxima, was in reality a member University, and Dr. M. W. Sanderson, Illinois of her new genus. The status of Joly's nymph Natural History Survey, for the loan of specimens has still not been established with certainty by of Neoephemero. Mr. G. S. Walley, Division of European workers nor, as far as I can determine, Entomology, Ottawa, kindly provided me with a has the species again been recorded. paratype of N. bicolor. All illustrations were Because of similarities in appearance of the prepared by Miss Esther Coogle, Staff Artist, nymph of Oreionthus and that of C aenis and Department of Biology, University of Florida. certain resemblances of the adults, Traver (1932) placed the former genus in the Baetinae, but Genus Neoephemera McDunnough indicated that the wing venation of the adult is Genotype: N. bicolor McD., Laprairie, Quebec, July 9, similar to that of the Ephemerinae. In this 1924. No. 1292, Canadian National Collection. Joly (1870, p. 142)-Caenis. 1This investigation was supported in part by a re­ Eaton (1881, p. 196)-Caenis. search grant, No. G-4058, from the National Institutes Eaton (1883, p. 140)-Tricorythus (?). of Health, Public Health Service. Lestage (1917, p. 373)-? Tricorythus. Date of acceptance, Oct. 4, 1954. McDtmnough (1925, p. 168)-Neoephemera. 33 34 Annals Entomological Society of America [Vol. 49

'fraver (1931, p. 103; 1932, p. 111)-0reianthus. youngi in lacking the srr1al1, dorsa1ly-projecting, Ulmer (1933, p. 199)-Neoephemera. median spine on tergite 2. Needham, 'fraver, and Hsu (1935, p. 289; p. 290)- N eoephemera; Oreianthus. MALE HoLOTYPE: Body length 9.8 mm.; meso­ Traver (1937, p. 34)-0reianthus. thoracic wings 8.U mm.; caudal filaments in­ Ulmer (1939, p. 484; p. 606)-Neoephemera; Oreianthus. complete. Berner (1950, p. 98)-0reianthus. Burks (1953, p. 42)-0reianthus=Neoephemera. Head: Eyes large, separated by a distance Berner (1953, p. 145)-Neoephemera. equal to about one-half the width of one of the compound eyes. \lertex pale; geminate, deep­ Wings hyaline; fore wings with basal costal brown lines extending from posterior margin of crossveins weak or vestigal; stigmatic crossveins lateral ocelli to occiput. Margin of occiput dark partly anastomosed; vein M2 curved toward vein brown. Frontal carina brownish; dark brown Cu1 near the wing base; vein A1 with from one to marking at its distal end. Heavy brown hori­ three crossveins attaching it to the hind margin zontal marking below the antennae extending (figs. I 9, 20). Hind wing with an acute costal almost from the median carina laterally to the projection (fig. 18). Legs normal; claws dis­ compound eyes. Antennae pale, marked with similar on legs. Pleural folds of abdominal deep-brown, longitudinal line running length of segments expanded laterally. Penis short, dome­ second segment; flagellum with a slight brownish shaped; separated by a V-shaped median cleft tinge. Occlli brownish; basally with usual dark on apical margin. Forceps with four segments, trarkings the two distal ones short (fig. 15). Three fully Thorax: Pronotum blackish brown around developed caudal filaments in both sexes. margins; very dark mid line; lateral portions Nymphs (fig. 9) grossly resemble Caenis but slightly paler than the margins. Mesonotum have two pairs of wing pads. Head almost much paler in color than the pronotum; blackish twice as wide as long. Second segment of penciling around the margins of the scutum; antennae covered with short, plumose hairs which scutellum pale in the median area and with are also present sparsely on vertex of head; blackish brown margins. Metanotum dark me­ such hairs are also present on the legs and abdo­ dially, paler laterally. Pleurae pale with dark, men. Prothorax longer at margins than at mid purplish-brown markings at the bases of the line, and margins may be considerably elongated wings and legs. Sternum blackish brown but at anterolateral angles (fig. 5); lateral margins mottled with pale spots between the bases of developed into thin expanded areas. Body each pair of legs; laterally brown. Prosternal widest at greatest width of mesothorax. Sub­ tubercle blackish brown and projecting ventrally median paired tubercules may be present at almost as far as the distal end of coxae. Legs. anterior margin . of pronotum and rnesonoturn Fore coxa pale with blackish-brown spot on outer (fig. 4); short, posteriorly-projecting median side; trochantcr pale with purplish-brown pen­ tubercule on rnetanotum. Legs long and spider­ ciling on outer surface; femur colored with intense like; claws simple, curved (fig. 11). Abdominal purplish gray markings distally and pale in segment I with a pair of vestigal, filamentous basal fifth; this basal pale area extending dis­ gills; gills of segment 2 fused to form elytroid tally along the front margin about one-half its plates covering the following four pairs (fig. 9). length; posterior margin colored deep purplish Prominent median spine on tcrgites 1 and 2; gray becoming much more extensive and cover­ tergites 2 to 8 may have a median carina which ing remainder of femur. Tibia colored much terminates in a short spine. Lateral margins of the same as the femur, pale basally except for a segments 3 to 9 prolonged posteriorly into short, longitudinal dark dash at the base; distal flattened processes that may be decurved (fig.4); four-fifths purplish gray, except at the extreme these processes exceptionally well developed on end where it becomes pale once more. Tarsal segments G to 9. Three caudal filaments with segments 1 to 4 pale but washed with purplish rings of stiff hairs at the joints. gray, fourth dark at distal end, fifth segment much deeper in color with tendency to form a band in middle portion; claws shaded with Neoephemera compressa n. sp. purplish gray; first tarsal segment very short, The adults of this new species can be separated about one-third the length of fifth, second and from the others of the genus by size and vena-. third equal, fourth slightly shorter than third, tional characters or bv color differences. N. and fifth slightly shorter than fourth. Middle compressa is distinct frOm the other two species and hind legs colored very much as fore legs in the size range in which it falls, N. bicolor and except for the fact that their coloration is even Jll. youngi, in the coloration of its legs and the more intense. Pale area of femur less than that absence of annulations on the caudal filaments. of fore femur; tibiae with a short dark dash at From IV. purpurea, it rray be separated by its the base on the inner side. Tarsal segments smaller size and by the presence in the fore wing with a tendency to form purplish rings on seg­ of only a single crossvein attaching A1 to the ments 2 to 5; fifth almost entirely purplish gray; hind margin. f\T. compressa also differs from l\7• claws as on fore legs. U'ings: Hyaline and as 1956] Berner: Neoephemera in. North America 35

FIGS. 1-4 .. 5 mphs to FIGS. 5-8. nymphs Obli4uc vte\ys. to fbercu1es half-gro\vn and nabdom Y inal .· w of thorax of ma tur~5 N. pur- h thoracic u . tubercules7 bicolor, showingDors.~l v~~oracic N compressa. · N. s ?w N. youngi. purea. 6. . · spines. purpurea. 2 · 8_ N. youngi. 31._ Neoephemera4N. bicolor. · N. compressa. 36 Annals Ent-Omological Society of America [Vol. 49 shown in figs. 22 and 23. All longitudinal veins paler at the proximal end. Tibiae of middle and pale; stigmatic area cloudy, crossveins anasto­ hind legs with small pale area basally, remainder mosed; single crossvein attaches Ai to hind deep purplish gray. Tarsal segments as in male, margin. Hind wing with a slight purplish-gray with color becoming more intense distally; first coloration at extreme base; costal projection so tarsal segment of middle and hind legs almost placed that it overlaps the anal crossvein probably completely pale, washed with light purplish providing a coupling device for the two wings coloration. when the insect is in flight. Wings: Hyaline, some slight, purplish shading Abdomen: Purplish brown dorsally; pale me­ near the base; costal border with a faint purplish dian line running the length of tergites 1 to 9; tinge; costa, subcosta and R, with purplish tenth completely dark. On tergite 2, the pale shadings, other longitudinal veins pale. Stig­ area expanding somewhat so that much of the matic veins weak and anastomosed. median section is occupied by a pale area; Abdomen: Almost uniformly purplish brown on tergites 3 to 5 the median line expanding at with a faint indication of a pale median line the anterior margin into a broad pale area beginning at the posterior portion of tergite 6 that runs along the anterior margin almost to and running full length of 7 to 9; tergite 10 the lateral line; on tergite 6 the expansion also lacking this pale median line. Ventrally, sternites occuring at the posterior margin and running mottled with purplish coloration; a clear spot pres­ laterally almost to the lateral border. Tergites ent at the anteromedian margin of segments 2 to 3 to 9 with small lateral, clear spots toward the 7. Lateral flanges deep purplish brown; caudal posterior margin. Spiracular areas of segments filaments washed with a purplish tinge. 1 to 9 deeply colored with blackish brown. On NYMPH: Body length 9 mm.; caudal filaments the tergites in which the lateral margins are 8.5mm. expanded, particularly 7 to 9, purplish brown Head: Vertex with numerous course hairs over coloration following the expansion; tenth tergite the surface, some present on the medial surface more or less uniformly colored with mottled of compound eyes. Hairs enlarged and frayed purplish-brown; some intensification of color in distally. Head crossed by purplish-brown band the median line. Posterior margins of tergites 8 between the eyes; vertex brownish with two sub­ and 9 more deeply colored, pattern forming a median clear spots; pale between ocelli. Ventrad blackish-brown band. Sternites much paler than to the antennae, between the eye and antenna! tergites with a slight wash of purplish gray in base, a deep-brown l:iand extending across the the median area of 6 to 9 and laterally on 1 to 5. face. Basal segments of the antennae with the At the anteromedial line of the sternites, a deep same type of course hairs that are present on blackish-brown mark which extends posteriorly the surface of the head; these hairs especially as a streak on sternites 6 to 9; laterally on each of numerous on the second antenna} segment where the sternites just below the dark marks of the they give it a very rough appearance; flagellum spiracular areas, a deep, purplish-black mark that pale and long. becomes progressively smaller posteriorly. Geni­ Thorax: Lateral margin of pronotum as shown talia pale; as shown in fig. 16. Caudal filaments in fig. 6; most of the pronotum deep brown with pale; basal segments with a slight brownish flanges pale; pair of very prominent submedian coloration. tubercles at the anterior margin of pronotum. FEMALE ALLOTYPE: Wing length 10 mm., Mesonotum with anterolateral corners expanded body 9 mm. (abdomen of specimen in somewhat into almost spine-like projections; a pair of contracted state); caudal filaments 10 mm. minute submedian tubercules at anterior mar­ Head: Eyes widely separated. Vertex mottled; gin; a median spine present at the posterior a deep, blackish-brown band on the occiput margin between the wing pads. Mesonotum medial to the eyes runs across the head just brownish, anterolateral corners dark brown. above the median ocellus. Coloration of the Metanotum completely concealed by wing pads. head dark. Carina blackish brown in its distal Sternum mottled with purplish-brown; a ven­ portion. Coloration of head below antennae like trally projecting process on the sternum between that of male. Base of antenna pale, second each pair of legs, prominent on the younger segment marked as in male; flagellum brownish; nymphs; on mature nymphs the projections tips broken in allotype. clearly visible between the anterior and middle Thorax: Marked much as in male except that mesonotum considerably darker with mid line blackish brown. Pleural and sternal markings EXPLANATION OF FIGURES similar to those of male. Prosternal process FIG, 9. N. compressa, half-grown nymph. similar to that of male. FIG. 10. N. youngi, half-grown nymph. Legs: More intensely colored than in male with FIGS. 11-14. Right hind leg of mature nymphs. 11. N. pale areas smaller than those of male. Tarsal purpurea. 12. N. compressa. 13. N. youngi. 14. N. bi'color. segments appearing almost annulate in fore leg. FIGS. 15-17. Genitalia of male adults. 15. N. purpurea. Femora of middle and hind legs only slightly 16. N. compressa. 17. N. youngi. 1956] Berner: Neoephemera in North America 37

10

9

-.. _

15

16 38 Annals Entomological Society of America [Vol. 49 pairs of legs, but only a bump between the old nymphs. The posterolateral spines of the hind legs. A brown spot present on the sternum abdomen are one of the notable features of the medial to each leg. young nymph. They also show a median line Legs: Covered with same type of hairs as on the tergites which includes the posteromedial seen on head and antennae. All segments of spines on segments 6 to 9. This light line is leg rather densely covered with these hairs, inter­ sometimes very pronounced. Because of the spersed with long normal hairs on the femur and coloration of the posterolateral spines the ab­ tibia. All legs long and spindly; purplish; femora domen may often appear to be banded at the more intensely colored at distal end; at the juncture margins. of trochanter and femur, on the ventral side, a Holotype: Male imago (reared), preserved in I large brown spot overlapping both segments. alcohol. Hamilton Co., Florida, Withlacoochee I Tarsus slightly paler at its proximal end; claw River at Georgia-Florida state line, March 14, heavy and curved; pale except at tip where it 1954. Collected by L. Berner. In University becomes deep brov.rn in color. Claws in striking of Florida collections. contrast to the dark color of rest of the leg. ALLOTYPE: Female imago (reared), preserved Abdomen: Abdominal segments expanded later­ in alcohol. Sarne data as holotype. In Uni­ ally into long, posterolateral projections, becom­ versity of Florida collections. ing more conspicuous on posterior segments PARATYPEs: 1 male subirnago (partially shed); where they are decurved. The posterior spine of 4 females (reared). Sarne data as holotype. In each segment approximately half the length of University of Florida collections. that tergite, except that on 9 the spine equal to the length of the tergite. In older specimens OTHER LocALITY RECORDS: Florida, Withla­ the posterolateral spines clear at the outer coochee River at several localities, January 28, margin and at the tip, but with purplish-brown 1954, (C. D. Hynes); Santa Fe River at Camp coloration running into the posterior portion: O'Leno State Park, October 31, 1953 (C. D. anterolateral portion of the spines clear. The Hynes). Georgia, Jchawaynochaway Creek at lateral margins of the segments covered with the Ga. Hwy. 91, November 27, 1953 (C. D. Hynes same type of course frayed hairs present on the and L. Bemer); Withlacoochee River, 4 miles head. Tenth segment without postero-lateral west of Valdosta, March 13, 1954 (Hynes and spines. Tergites 1 and 2 with dorsally-project­ Berner). AB specimens in University of Florida ing, pale median spines between the wing pads. collections. The spine on segment 2 especially prominent and The nymphs of 1'l. compressa occur in streams also covered ,•;rith the course hairs. Tergites G of slow to moderate flow. The Withlacooehee to 8 free of gill covers, ¥.•ith posteriorly projecting River, where the nymphs were found in some spines at the rredian line; these spines much less abundance, is a deep, slow-flowing stream conspicuous than those on tergites 1 and 2 and bordered by ash, willow, cypress and black-gum not erect. Abdomen deeplv colored with pur­ trees. VVhere the trunks of these trees are sub­ plish brown on the dors'tm; ~rentra1ly paper with merged, they are usually covered by a densely mottled appearance. The whole venter covered growing species of moss. It was in this moss with coarse, fraved hairs. At the anterolateral that the nymphs were found, usually after a margin of each of the sternites, a purplish­ careful examination. They are slowly moving in­ brown dash directed inward, the dash being less sects which are sometimes difficult to see in the prominent but more elongate on the posterior dark vegetation and it is only by removing segments and obsolescent on stcrnite 9. Caudal large clumps of the plant and shaking them that filaments approximately equal in length; brownish the nymphs are dislodged. Many were removed and with stiff, coarse hairs at the joints. hy picking them from the moss after it was Younger nymphs are particularly striking in taken from the water. The nymphs ¥.'ere as­ appearance and the natr.e that I have given this sociated with those of Baetisca obesa, which were species is derived from the pronounced dorso­ very common in the moss. Amphipods and ventral flattening of the thorax. The tubcrcules isopods were also numerous in this association. of the anteromedian margin of the pronotum are Nymphs were often found in some abundance by very conspicuous 2 s is another pair at the an­ running a dip net through debris anchored in terior margin of the tr:iCsonotum. The single· flowing water. posterior tuberC'ulc of the rr.esonotum is also verv At the Ichawaynochaway Creek near Newton, prominent. As the nymphs mature, the thora~ Georgia, nymphs were taken by washing the swells in size so that it assutres a more normal submerged roots of plants which hang exposed shape and the anterior tubercles arc almost at the banks. A few nymphs were also collected obliterated. The heads of younger nymphs are from logs which had been in the stream for a often completely pale except directly behind the long period and had become very soft. The eyes, where they are deeply colored with purplish greatest number of nymphs from this stream were, brown. Legs of the young nymphs may also be however, taken from clumps of moss of the same paler basally instead of being wholly dark as in species as that growing. in the Withlacoochee 1956] Berner: Neoephemera in North America

25 FrGs. 18-25. Wings of adults. 18-19. N. purpurea. 20-21. N. bicolor. 22-23. N. compressa. 24-25. N. youngi. 40 Annals Entomological Society of America [Vol. 49

River. No nymphs were found in the swiftest of the fact that she failed to mention the pair of part of the stream. low submedian tubercles on the anterior margin Adults were reared from mature nymphs col­ of the pronotum. Traver also pointed out that lected from the Withlacoochee River in January the short median tubercle on the mesonotum be­ and March. The earliest emergence occurred t ween the wing pads was, in the original descrip­ in late February and the latest was recorded in tion, in error described as being on the metanotum. early April. The subimagal stage lasts approxi­ The nymph is by far the largest of any of the mately twenty-four hours. North American species of Neoephemera and is My collections indicate that N. compressa may readily distinguished from the others in the be a southern species restricted to streams of the mature form by its size. The lateral profile of Coastal Plain. the nymph is distinctive as indicated in the

FrG. 26. The Withlacoochee River near Valdosta, Georgia, March 21, 1954. Neoephemera compressa nymphs were taken from the moss growing on the submerged trunks of these trees and from debris anchored in the stream.

Neoephemera purpurea (Traver) drawing (fig. 1). The tubercules on the pro­ Traver (1931, p. 103)· Oreianth11s p11rp11re11s. Type in and mesonotum arc relatively short and incon­ Cornell University Collections. spicuous on the mature nymph as compared with Needham, Traver, Hsu (1935, p. 291)-0reianthus p11r­ p11re11s. those of bicolor and compressa. On half-grown Traver (1937, p. 34)-0reianth11s p11rp11re11s. nymphs the tubercules are exaggerated but are Burks (1953, p. 43 )-Neoephemera p11rpurea. still considerably less so than on nymphs of N. This species was described by Traver as compressa at a comparable stage of development. Oreiantlms purpureus in 19:31. In her paper she The specimens examined by Traver and re­ gave a very complete description of both female ported on in publications subsequent to the imago and the nymph; however, in this earlier original description have all represented mature description one error was made which was later nymphs. The appearance of a half-grown speci­ corrected in her 1937 report on the mayflies of men is shown in fig. 1. the southeast. The description is entirely ade­ When Traver (1937) collected the nymphs of quate and the 1937 correction simply makes note N. purpurea in North Carolina, she found them 1956] Berner: Neoephemera in North America 41 usually beneath small, isolated boulders or ir­ sembled those of N. compressa but differ in regularly-shaped sedimentary rocks in swift water, minor details as indicated in the key. Because or under the large, flat rocks that form the main of the presently known distribution of N. bicolor, bed of the stream. Here she found only few I believe that Dr. Burks identification is correct specimens, never taking more than two or three and I am calling these specimens by that specific nymphs from under a single rock. In 1948 while name. I was collecting in the North Fork of the French The'nymph of N. bicolor has never been de­ Broad River at Rosman, I found several half­ scribed. I shall not detail the characteristics of grown nymphs of this species living in debris the Michigan specimens as they are practieally in slow-flowing water. A search of the larger the same as those of N. compressa except in the rocks produced no specimens. During the sum­ following respects: the pronotal spines are more mer of 1954, I again had an opportunity to widely spaced and are slightly less prominent, collect mayflies in the mountain streams of North and the posterolateral abdominal spines are Carolina and at this time found nymphs of . slightly shorter and are less strongly curved N. purpurea in Deep Creek and in the N antahala ventrally. These details are shown in figs. 3, River living commonly in debris caught in flowing 4, 6, and 7. water. Where bushes had fallen into the stream Mr. G. S. Walley of the Division of Ento­ and had remained there for some time, the mology, Ottawa, has kindly furnished me with a nymphs could be taken by dislodging the debris paratype and a topotype of N. bicolor. A com­ to which they were attached. In the N antahala, parison of these specimens with the three southern as many as seventeen nymphs were taken at one species has made it possible for me more clearly time by gathering up clumps of exposed, clean­ to delineate the forms in the key. wa.shed roots of vegetation in shallow water. Specimens examined: Quebec, Laprairie, July The nymphs clung tenaciously but could be 9, 1924 (G. S. Walley), paratype; July 7, 1925 shaken or picked loose. A few nymphs were (F. P. Ide). Michigan, Riffle River, Omer, May also collected by lifting large stones where the 21, 1936 (Frison and Ross)-nymphs. flow was swift and holding a screen downstream; There was no ecological information ac­ however, this method of collecting was much companying the nymphs of N. bicolor that were more tedious and less productive than working collected in Michigan. Because of their close re­ in debris and roots. semblance to the nymphs of N. youngi and com­ Distributional records indicate that this species pressa, I believe that they occupy similar habitats, is confined to the swift streams of the Appalachian occurring in rivers and large streams that have a mountains. slow to moderate flow. Specimens examined: North Carolina, David­ son River, Pisgah Forest, (J. R. Traver), June N eoephemera youngi Berner 2 19, 22, 1929, C.U. ; June 30, 1930 (Holotype) Traver (1937, p. 83)-0reianthus sp. No. 1. C.U.; June 12-22, 1936, C.U.; North Fork of Bemer (1950, p. 99)-0reianthus sp. No. 1. French Broad River at Rosman, August 19, 1948 Bemer (1953, p. 145)-Neoephemera youngi. Type de­ 2 posited in University of Florida Collections. (L. Berner) U.F. ; Deep Creek, Bryson City, August 12, 1954 (L. Berner) U.F.; Nantahala Among the specimens of N eoephemera in the River. August 16, 1954 (L. Berner) U.F. Ten­ collection of Illinois Natural History Survey, nessee, Elkmont. October 10, 1940 (L. Giovan­ there is a single, adult male from Echeconnee noli) U.F. Georgia, Neel Gap, June 8, 1945 Creek, Georgia, identified by Dr. B. D. Burks as (P. W. Fattig) I.N.H.S.'; Rock Creek, May 14, N. bicolor. A careful examination of this speci­ 1931 (P. W. Fattig), C. U.; Satolah, May 4, men shows that it is in reality N. youngi. Ap­ 1929 (J. G. Needham) C.U. Virginia, Piney parently on the basis of this male, Dr. Burks River, August 1, 1953 (J. Pugh) U.F. West (1953) reported that N. bicolor occurs in Georgia, Virginia, Elk Garden, August 21, 1930, C. U. a distribution which would have been unlikely as it is known elsewhere only from Michigan and N eoephemera bicolor McDunnough Quebec. Further, Dr. Burks reported that N. McDunnough (1925, p. 168)-Neoephemera bicolor. 'l'ype purpurea occurs in Florida; intensive studies of in Canadian National Collections. Needham, 1'raver, Hsu (1935, p. 289)-N. bicolor. the Ephemeroptera of this state have never pro­ duced a nymph of purpurea nor have I ever Through the courtesy of the Illinois Natural taken the species south of the Appalachians. It History Survey, I was privileged to study their is my belief that N. youngi is essentially a costal collection of N eoepkemera. Included in the lot plain species that may be much more widely of specimens was a small series of n:yn1phs from distributed through this physiographic province Michigan identified by Dr. B. D. Burks as than is indicated by specimens in collections. ?N eoepkemera bicolor McD? These nymphs re- N. youngi is closely related to bicolor and compressa and with these two species forms a 2C. U.-Cornell University collections; U. F.­ University of Florida collections; I.N.H.S.-Illinois clearly distinct unit of the genus N eoepkemera. Natural History Survey collections. Size, venation, and characters of the immatures 42 Annals Entomological Society of America [Vol. 49 reveal this grouping very clearly as indicated in 11• Wings 8 to 10.5 mm. in length; veins, except those the keys to adults and nymphs. of costal border, pale. A single crossvein attach- ing A1 to hind margin {fig. 20). 2 Distribution and specimens examined: Gener­ 2. Small purplish band present in distal four-fifths of erally distributed throughout northwest Florida femora; tarsal segments 2 to 4 of middle and hind from the Apalachicola River system westward. legs ringed with purplish-gray. Caudal filaments In Georgia from Decatur Co., Mosquito Creek, annulate ...... youngi 2'. Coloration of legs otherwise. Caudal filaments April 5, 1953, March 28, 1954 (L. Bemer) U.F.; without annulations...... 3 Town Creek, May 14, 1931 (P. W. Fattig) 3. Femora usually pale; tarsi smoky, not annulate. C.U.; Oconee River near Greensboro, April 9, Each stemite with lateral black dot...... bicolor 3 1• Fore femur extensively colored with purplish­ 1931 (P. W. Fattig) C.U.; Upatoi Creek near gray; tarsus purplish-gray, fifth segment with Columbus, April 29, 1931 (P. W. Fattig) C.U.; purplish-gray band. Other legs heavily shaded Echeconnee Creek, 11 miles south of Macon, with deep purplish gray, tarsal segments annulate. May 5, 1939 (P. W. Fattig) LN.H.S. South Sternites with lateral black dot as well as a blackish- Carolina, North Edisto River at Orangeburg, brown dot at anteromedial line ...... compressa October 21, 1952 (H. P. Nicholson) U.F. The habitats occupied by nymphs of N. youngi REFERENCES CITED were discussed in some detail in "The Mayflies Bemer, Lewis. 1950. The mayflies of Florida. Univ. of Florida" (Berner, 1950) and will not be rede­ Fla. Studies, Biol. Sc1. Ser. 4 (4): i-xii, 1-267. 88 scribed here. In many respects they are similar figs., 24 pls., 19 maps. N. compressa 1953. New records from Florida and a descrip­ to those from which was taken, tion of a new species. Fla. Ent. 36 (4): 145-9. 3 figs. although no youngi has ever been taken from Burks, B. D. 1953. The mayflies, or Ephemeroptera, of moss around the trunks of submerged trees. Illinois. Bull. Ill. Nat. Hist. Surv. V. 26 (1): 1-216, KEY TO NYMPHS 395 figs. Eaton, A. E. 1881. An announcement of new genera of 1. Body length (head to tails) 14-17 mm. Antero­ Ephemeridae. Ent. Mo. Mag. 17: 191-7; 18: 21-7. lateral projections of pronotum strongly produced 1883-1888. A revisional monograph of recent Ephem­ (fig. 5); anterolateral projections of mesonotum eridae or mayflies. Trans. Linn. Soc. London, Sec. prominent and rounded...... purpurea Ser. Zool. 3: 1-352. 65 pls. 11 Body length 8-10 mm.; anterolateral corners of pro~ notum only slightly elongated (fig. 6)...... 2 Joly, Emile. 1870. Contributions pour servir a l'histoire 2. Posterolateral abdominal spines strongly produced naturelle des Ephemerines. Bull. Soc. d'Hist. Nat. (figs. 4, 9). Median spines present at posterior de Toulouse 4: 142-50. Pl. 3. margins of abdominal segments 6-8 (fig. 4). An­ Lestage, J. A. 1917. Contribution a l'etude des larves terolateral comers of mesonotum pointed (fig. 6); des EphemCre.<> palearctiques. Ann. Biol. Lacustre 8: pronotal submedian tubercules prominent (fig. 6). 3 213-457. 54 figs. 2.' Posterolateral abdominal spines only moderately McDunnough, J. 1925. New Canadian Ephemeridae produced (fig. 2, 10). No median spines on pos­ with notes, III. Can. Ent. 57: 168-76. Pl. 4. terior margins of abdominal segments 7 and 8 and only faintly indicated on 6 (fig. 10). Anterolateral Needham, J. G., Jay R. Traver, and Yin-Chi Hsu. 1935. comers of mesonotum rounded (fig. 8); pronotal The biology of mayflies. Pp. xvi +759. 168 figs., submedian spines much reduced in mature nymphs 40 pis. Comstock Publishing Co., Ithaca. (fig. 8) ...... youngi Traver, Jay R. 1931. A new mayfly genus from North 3. Posterolateral abdominal spines strongly produced Carolina. Can. Ent. 63: 103-9. PL 7. and curved ventrally (fig. 4). Pronotal submedian 1932. Mayflies of North Carolina. Jour. Elisha spines only slightly separated (fig. 6). . . . . compressa Mitchell Sci. Soc. 47: 85-161, 163-236. Pls. 5-12. 3.' Postero-lateral abdominal spines less strongly 1937. Notes on mayflies of the southeastern states produced and less strongly curved ventrally (fig. 3). (Ephemeroptera). Jour. Elisha Mitchell Sci. Soc. Pronotal submedian spines widely separated (fig. 7) 53: 27-86. PL 6...... bicolor Ulmer, George. 1933. Aquatic of China. Art. VI. Revised key to the genera of Ephemeroptera. KEY TO ADULTS Peking Nat. Hist. Bull. 7: 195-218. 2 Pls. 1. Wings 13.5 to 17 mm. in length; longitudinal veins 1939-1940. Eintagsfleigen (Ephemeropteren) von den of fore wings purplish black; two crossveins at­ Sunda-Inseln. Archiv ft1r Hydrobiol. Suppl. Band, taching A1 to hind margin (figs. 18, 19) ..... purpurea 16: 443-580, 23 Pls.