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The Mayflies of North and Central America

The Mayflies of North and Central America

George F. Edmunds
Steven L. Jensen
Lewis Berner
Copyright Date: 1976
Edition: NED - New edition
Pages: 344
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  • Book Info
    The Mayflies of North and Central America
    Book Description:

    Mayflies have fascinated man for centuries because of the brief span of their adult lives. These aquatic insects spend most of their lives as nymphs in water, then develop into winged stages and soon die, most species having an adult life of only two or three days. This brevity is implied in the very name of the order, Ephemeroptera. The mayflies are almost worldwide in distribution, being found everywhere except in Antarctica, the extreme Arctic, and many small oceanic islands. All by three of the twenty families in the world occur in North or Central America, the regions covered in this volume. The book provides a modern, useful, and well-illustrated key to the adults and nymphs. Data on habitats, behavior, and life history are given for each genus. Characteristics of nymphs and adults are given for families, subfamilies, and genera, with brief accounts for extralimital families. A discussion of methods of collecting and preserving specimens precedes the main portion of the text. The book is generously illustrated with drawings, photographs, and a map. The role of aquatic insects as indicators of water pollution has received increasing attention, and in this connection this book will be of special interest to those concerned with pollution problems. Mayflies, besides indicating the presence of pollutants, also help remove such substances from the waters, the authors explain. As a basic reference work, the book is essential for all biological science libraries. Many fly-fishermen are amateur students of mayflies, since the nymphs of larger species are used as bait. With the help of this volume the fisherman can acquire a greater knowledge of aquatic entomology and relate to his sport.

    eISBN: 978-0-8166-6225-8
    Subjects: Zoology

Table of Contents

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  1. Front Matter
    (pp. i-vi)
  2. Table of Contents
    (pp. vii-viii)
  3. Preface
    (pp. ix-2)
  4. The Biological Role of Mayflies
    (pp. 3-48)

    The mayflies have fascinated man for centuries because of the brief span of their adult lives. These aquatic insects spend most of their lives as nymphs in water, then develop into winged stages and soon die. The white mayfly,Ephoron album, has a winged stage lasting a bare ninety minutes. In fact, the female of this species lays her eggs and dies as a subimago, that is, before ever achieving full adulthood. Similar brevity is almost certainly the case with several other species. Although some species survive as adults for ten days and may live for weeks, most mayflies have...

  5. Key to Nymphs
    (pp. 51-83)

    1 Thoracic notum enlarged to form a shield extended to abdominal segment 6, gills enclosed beneath shield (fig. 26); nymph shown (fig. 423); mid-boreal, widely distributed in eastern and central North America, rare in west....Baetiscidae,Baetisca(p. 269)

    Thoracic notum not enlarged as above; abdominal gills exposed ...................................................2

    2(1) Mandibles with large tusks (t) projected forward and visible from above head (figs. 30-34) ..............................3

    Mandibles without such tusks (figs. 42, 53, 56, 135-138) . . .9

    3(2) Gills on abdominal segments 2-7 forked, without margins fringed (fig. 29); mid-boreal, western North America south to California and Utah ............................................Leptophlebiidae,Paraleptophlebia, in...

  6. Key to Adults
    (pp. 84-122)

    NOTE: Unless specified otherwise, the venation referred to in the couplets is that of the forewing.

    1 Wing venation greatly reduced, apparently only three or four longitudinal veins behind R₁ (figs. 195a, 196a); body black .......................................Oligoneuriidae, 41

    Wing venation complete or only moderately reduced, numerous longitudinal veins present behind R₁ (figs. 197-201, 217-224); body color variable ...................................2

    2(1) Penes of male longer than forceps (fig. 204); antennae of female inserted on prominent anterolateral projections (fig. 202); four or more long cubital intercalaries usually present (fig. 197a); rare; mid-boreal, southeastern United States south to Florida Behningiidae,Dolania(p. 274) Penes of...

  7. The Families of Mayflies
    (pp. 123-129)

    In the familial accounts distinctive taxonomic characters are given for each family, the superfamily assignment is noted, and the most probable cladistic relationships are indicated. For each family and subfamily the total world distribution is discussed. Familial and subfamilial diagnostic characters are given for nymphs and adults of each family and subfamily. Brief accounts are given for the three families not found in the Americas. Within each subfamily the discussions of North and Central American genera and subgenera cover nymphal characteristics, nymphal habits and habitats, life history, adult characteristics, mating flights, taxonomy, and distribution. Reference is made to all nymphal...

    (pp. 130-147)

    The Siphlonuridae of the superfamily Heptagenioidea exhibit more primitive characters than any other extant family of mayflies. The family appears to be an assemblage of forms from an early adaptive radiation in the order. Most of the families of Heptagenioidea can be traced from the Siphlonuridae, and some of the characters of the south temperate subfamily Oniscigastrinae suggest that it is near the base of the phyletic line that leads to the nonheptagenioid families.

    Most of the genera of the family are adapted to cool waters and are most diverse in the Holarctic region and in Australia, New Zealand, and...

    (pp. 148-151)

    This small family of Heptagenioidea appears to be a derivative of the protosiphlonurine Siphlonuridae rather than a relative of the genusAmetropus. The two included genera are very closely allied, although the adults differ quite markedly. The nymphs ofMetretopusare more delicate, softer, and more slender. The two genera are not easily separated as nymphs, partly because the nymphs of all of theSiphloplectonare not known and partly because the known species have diverse gill types. However, the keys should allow for discrimination of the two genera. The adults are readily distinguished by the venation of the forewings....

    (pp. 151-151)

    This family is recognized for a single New Zealand species,Siphlaenigma janaePenniket. It is clearly intermediate between the Baetidae and Siphlonuridae. The nymphal behavior is quite similar to that of other stream Baetidae, but the abdominal ganglia and Malpighian tubules are siphlonurine. The placement of a single species in a separate family is a questionable practice, but it must be weighed against the difficulty of defining the Baetidae or Siphlonuridae ifSiphlaenigmais included in either family. To be consistent with classificatory practice in the mayflies, the taxon might better be placed as a subfamily of the Siphlonuridae.


    (pp. 152-154)

    The Ametropodidae appear to be Heptagenioidea that diverged from the ancestral stock early and now constitute an isolated group. The family is recognized for the single genusAmetropus. The various other genera that have been placed in the family are now assigned to Pseudironinae of the Heptageniidae, Acanthametropodinae, and Siphlonurinae of the Siphlonuridae or to the Metretopodidae. Long claws are primarily an adaptation for sand dwelling, and this adaptation seems to influence the proportions of other parts of the leg in adults and nymphs. These characters have been used to justify the placement of genera of diverse origins in the...

  12. Family BAETIDAE
    (pp. 154-181)

    This family is a member of the Heptagenioidea. The Baetidae almost certainly have been derived from proto-Metamonius-complex siphlonurine Siphlonuridae of Chile, Argentina, Australia, and New Zealand. The New Zealand genusSiphlaenigma(Siphlaenigmatidae) is almost perfectly intermediate between the Siphlonuridae and the Baetidae and must have been derived from an early pre-baetid. In North America the Siphlonurinae most closely related and most similar to the Baetidae, especially as nymphs, are members of the genusAmeletus.

    The family is widespread, being found on all continents and on many islands. It is absent from New Zealand, although its closest relative,Siphlaenigma, occurs there....

    (pp. 181-185)

    This family of Heptagenioidea is almost certainly derived from proto-isonychiineSiphlonuridae. The adults of this family are among the most distinctive mayflies known. The nymphs are also readily recognizable, having diverged considerably from their nearest relatives. The Asian genusChromarcys, the most primitive member of the family, is placed in a separate subfamily Chromarcyinae or with fossil genera in the Hexagenitinae, but all other genera are in the subfamily Oligoneuriinae.

    Although predominantly pantropical, the Oligoneuriinae extend north in the Old World to Central Europe, Afghanistan, and Japan and in the New World to Saskatchewan. To the south they extend to...

    (pp. 186-213)

    The Heptageniidae were almost certainly derived from near the base of the phyletic line which evolved also intoIsonychia. The nymphs of Arthropleinae have specialized mouthparts, but the adults and nymphs also show many primitive characters; therefore, the genusArthropleais sometimes placed in a separate family.

    The family is diverse and abundant in the Holarctic, especially in the northern regions. A few genera are found in the Ethiopian region, these being closely allied to the more diverse Oriental fauna which extends to the Sunda Islands. The family is absent from the Australian region (the one published record from this...

    (pp. 214-239)

    This family is placed in the Leptophlebioidea. The family is relatively isolated and retains a number of primitive characteristics. It shares a number of characters with Ephemerellidae and Tricorythidae and also with the Ephemeroidea. The pre-leptophlebiid mayflies may have given rise to all of the families except those assigned to the Heptagenioidea. For this reason, the decision about which other families should be in the Leptophlebioidea, if any, is not an easy one to resolve to the satisfaction of most workers. The Leptophlebiidae seem to occupy a postgroup relationship to the ancestral form which also gave rise to the Ephemerellidae...

    (pp. 239-252)

    The Ephemerellidae probably arose from the phyletic line leading to the Leptophlebiidae and the Ephemeroidea. The most closely related extant mayflies are the Tricorythidae, which appear to be derived from Ephemerellidae. The Ephemerellidae and the Tricorythidae of North and Central America do not indicate this relationship as well as do the African genera of the two families. The family Ephemerellidae is assigned to the Leptophlebioidea, but because the Leptophlebiidae appear to have given rise to the Ephemeroidea, some workers prefer to isolate the Ephemerellidae and the Tricorythidae in a separate superfamily, Ephemerelloidea.

    The family is widespread on the continents, but...

    (pp. 253-260)

    This family has many specialized characters and appears to have arisen from proto-teloganodine Ephemerellidae; it is usually assigned with the Ephemerellidae to the Leptophlebioidea. The family is extremely diverse in Africa, where all the subfamilies (Tricorythinae, Ephemerythinae, Machadorhythinae, Dicercomyzinae, and Leptohyphinae) are present. The Tricorythinae extend through the Oriental region. Leptohyphinae is the only subfamily occurring in the Americas.

    This subfamily reaches its maximum diversity in South America. The two North American genera and the one Central American genus are northward extensions of the Neotropical fauna in whichLeptohyphesandTricorythodesare each represented by many species; one additional species...

    (pp. 260-263)

    This family is most closely allied to Caenidae, and the proto-Neoephemeridae were ancestral to the caenids. The two families constitute the Caenoidea, which in turn are allied closely to the Prosopistomatoidea.

    The nymphs of Neoephemeridae are distinguishable from those of the Caenidae by only a few minor external characters. The rounded lobe on the anterolateral corners of the mesonotum which distinguishes North American nymphs of Neoephemeridae from those of Caenidae is not present in all Neoephemeridae, but all Neoephemeridae have hind wing pads and the fibrilliform portion of the gill on segments 3-6. The adults superficially resemble the Ephemeroidea, especially...

  19. Family CAENIDAE
    (pp. 263-269)

    This family of Caenoidea has a number of specialized characters in both adults and nymphs. The family is almost certainly derived from the proto-Neoephemeridae of thePotamanthellusgroup lineage. The nymphs of Caenidae are so similar to some Neoephemeridae that only the absence of hind wing pads and the absence of the fibrilliform tuft on the gills of segments 3-6 serve to distinguish the nymphs of Caenidae. The adults of Caenidae, however, are remarkably different from the ancestral pattern which persists in the Neoephemeridae.

    The family occurs in a variety of ponds, lakes, streams, and rivers over most of the...

  20. Family BAETISCIDAE
    (pp. 269-272)

    The family Baetiscidae contains only the genusBaetisca. The family is an isolated one whose nearest relative is the Old World genusProsopistoma, the only genus of Prosopistomatidae. The two families are placed in the Prosopistomatoidea, a superfamily believed to have been derived from ancestral pre-Caenoidea.

    The family is endemic in North America. SeeBaetiscafor characteristics and distribution.

    Nymphal Characteristics. Body length: 4-14 mm. Body stout. Head frequently with prominent frontal projections and spinous projections from anterior of genae. Large mesonotal shield, or carapace, completely covering dorsum of mesothorax, metathorax, and abdominal terga 1-5 and basal half of tergum...

    (pp. 272-273)

    The Prosopistomatidae (superfamily Prosopistomatoidea) and Baetiscidae probably arose from a common ancestor that was a pre-caenoid. The Caenoidea constitute another branch of this phyletic line. It seems unlikely that the highly specialized carapace of the Prosopistomatoidea nymphs would have evolved from nymphs which already have a modification for gill protection in the form of the specialized quadrate gill 2 of the Caenoidea.

    The family is recognized for a single genus that is widespread in the Old World tropics, with one species in Europe. The genus is found in Africa, Madagascar, Ceylon, the Sunda Shelf, the Philippines, New Guinea, and Europe....

    (pp. 273-276)

    This family is an isolated and primitive phyletic line within the superfamily Ephemeroidea. It seems unlikely that the Behningiidae could have evolved from any of the true burrowing nymphs that have the forelegs modified for digging or even from the nonburrowing Euthyplociidae and Potamanthidae with large mandibular tusks. The ancestral precursor of the Behningiidae must have had a nymph in which the tusks were absent or small, the forelegs were not modified for burrowing, and the gills on segment 1 were single. The ancestral nymph could have resembled the EuropeanPotamanthus luteusLinnaeus. The nymphs are unique among Ephemeroidea in...

    (pp. 276-278)

    Potamanthidae is one of the most primitive families of the Ephemeroidea, a superfamily most probably derived from leptophlebiidlike ancestors. The family is most diverse in the Oriental realm and in Palearctic Asia. The genusPotamanthus, which is the only genus of the family in North America, is widely distributed in the Holarctic. See genusPotamanthusfor characteristics and distribution in North America.

    Nymphal Characteristics. Body length: 8-15 mm. Nymph of sprawling type. Frontal margin of head somewhat rounded. Mandibular tusks projecting forward and visible from above; with only inconspicuous setae and short spines on dorsal surface and lateral margins. Femora...

    (pp. 278-281)

    This family of Ephemeroidea has relatively unspecialized characters, the nymphs lacking specializations for burrowing and the adults with abundant venation. The family is probably a derivative of pre-potamanthid Ephemeroidea.

    The family is of pantropical distribution with three of the seven genera occurring in South America. Of these, two genera are found in Central America and southern Mexico.

    Nymphal Characteristics. Mandibular tusks longer than head and with numerous long hairs. Femora flat and broad, tibiae and tarsi cylindrical, not modified for burrowing. Gills on abdominal segment 1 small and ovoid. Gills on segments 2-6 extend laterally from abdomen, fringes longer than...

  25. Family EPHEMERIDAE
    (pp. 281-294)

    The Ephemeridae are Ephemeroidea that probably arose from prepotamanthid ancestors. The family includes two subfamilies. The Old World Palingeniidae presumably were derived from proto-Pentageniaof the Ephemeridae. McCafferty (1972) has placedPentageniain a separate family, Pentageniidae. We have recognized this proposed change in the classification to the extent of placingPentageniain a separate subfamily.Pentagenianymphs seem to be at the palingeniid grade of evolution, but the adults remain undifferentiated from the ephemerids. Comparisons ofPentageniawith primitive members of the Palingeniidae are needed to clarify the taxonomic relationships in this group. It may be that the Pentageniinae...

    (pp. 294-294)

    The Palingeniidae are Ephemeroidea that are clearly derived from a proto-Pentageniaancestor. The nymphs ofPentageniahave in fact evolved to the palingeniid grade, but the adults have not diverged from the characters of the Ephemeridae. The adults of the Palingeniidae are strongly differentiated from the Ephemeridae, includingPentagenia.

    The Palingeniidae are widespread in the Old World, where they are found in the large lakes and rivers of Eurasia, New Guinea, Madagascar, and questionably Africa. Except forPalingenia, the family is rather poorly known. It is not known to occur in the Americas, but this will change if further studies...

    (pp. 295-302)

    The somewhat diverse Polymitarcyidae, whose nymphs are specialized for burrowing, were probably derived from the Euthyplociidae, a family whose nymphs lack specializations for burrowing.

    The family is widespread on the continents but absent from the Australasian realm. There are three subfamilies of which two occur in North America. The Asthenopodinae are pantropical and are found in South America, Africa, and the Oriental region. Most species are pale milky white with darker markings, and all seem to have crepuscular to nocturnal mass mating flights.

    The family and subfamily names Polymitarcyidae and Polymitarcyinae are reserved for these taxa under rule 40 of...

  28. Appendix I A Guide to Changes in Scientific Names
    (pp. 305-308)
  29. Appendix II Suppliers of Collecting and Rearing Equipment
    (pp. 309-310)
  30. Bibliography
    (pp. 313-322)
  31. Index
    (pp. 325-330)