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The family Baetidae (InsectaEphemeroptera) synthesis and futurechallengesJean-Luc Gattolliat a amp Carolina Nieto ba Museum of Zoology Place de la Riponne 6 LausanneSwitzerlandb CONICET-INSUE Fac de Cs Naturales e IML TucumaacutenArgentinaPublished online 24 Nov 2009

To cite this article Jean-Luc Gattolliat amp Carolina Nieto (2009) The family Baetidae (InsectaEphemeroptera) synthesis and future challenges Aquatic Insects International Journal ofFreshwater Entomology 31sup1 41-62 DOI 10108001650420902812214

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The family Baetidae (Insecta Ephemeroptera) synthesis and future

challenges

Jean-Luc Gattolliata and Carolina Nietob

aMuseum of Zoology Place de la Riponne 6 Lausanne Switzerland bCONICET-INSUE Facde Cs Naturales e IML Tucuman Argentina

(Received 19 December 2009 final version received 10 February 2009)

The systematics of the Baetidae has been the subject of much attention during thelast three decades with descriptions of new species and genera as well as severalgeneric revisions The family now encompasses about 100 genera and 900 specieswhich constitute one-quarter of the worldrsquos mayfly diversity It is thus anopportune time to evaluate the pertinence of these works The diversity of theBaetidae of the different realms is discussed with emphasis on the rate ofendemism and biogeographic affinities We have also tried to identify thegeographical areas where we need more data The Afrotropics and Neotropicspossess the most diversified fauna with the highest degree of endemism Thefaunas of the Palaearctic and Nearctic realms are better known but areless diversified especially at a generic level Oriental and Australasian faunasare still poorly known but are expected to be diversified especially in the tropicalregions

Keywords Baetidae biogeographic realms diversity systematics

Introduction

In 1815 Leach divided the tribe Ephemerides into two families depending on thenumber of caudal filaments in the imaginal stage He established the family Baetidaefor species with two caudal filaments and the Ephemeridae for species with threefilaments In the Baetidae he established two genera Baetis for species with fourwings and Cloeon for species with two wings In the last 200 years the systematics ofthe Baetidae has greatly improved and this family now encompasses about 100genera and 900 species which constitute one-quarter of the worldrsquos mayfly diversityExcept for Antarctica and New Zealand Baetidae present a cosmopolitandistribution

Baetidae constitute a relatively homogenous family They are distinguished in theimaginal stage by the presence of turbinate eyes in the male imago (except in thesingle Neotropical genus Aturbina (Lugo-Ortiz and McCafferty 1996a)) detachedIMA and MA2 forewing veins the presence of simple or double free intercalaryveins in the forewing (partially or completely reduced in a few taxa) hind wing

Corresponding author Email jean-lucgattolliatvdch

Aquatic Insects

Vol 31 Supplement 1 2009 41ndash62

ISSN 0165-0424 printISSN 1744-4152 online

2009 Taylor amp Francis

DOI 10108001650420902812214

httpwwwinformaworldcom

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reduced or absent three-segmented mid and hind tarsi and membranous penis(Edmunds et al 1976) Larvae are also unique among mayflies in having the lateralbranches of the epicranial suture anterior to the lateral ocelli and the ventralorientation of the dorsal lobe at the apex of femora (Wang and McCafferty 1996)Larvae are pisciform they generally possess long antennae and simple or doubleovoid gills on segments IndashVII or IIndashVII

Baetidae are very common in any kind of freshwater they are mainly diversifiedin unpolluted running water especially in the tropics Although they are lessdiversified in standing waters with genera like Cloeon the Baetidae constitute animportant part of the insect biomass in ponds Most species of Baetidae arecollector-gatherers feeding mainly on detritus Unusual adaptations to peculiar dietsoccur in a few genera Twelve species belonging to nine genera are considered to becarnivorous They feed mainly on larvae of Diptera but some are specialised infeeding on other mayflies (Gattolliat and Sartori 2001 McCafferty and Sun 2005)Although these carnivorous genera present similar adaptations of the mouthpartsthey clearly belong to different lineages (Monaghan et al 2005) In several lineagesspecies or genera show adaptations for scrapping algae on the top of stones (Lugo-Ortiz and McCafferty 1997 Gattolliat 2001b) Some genera have an advancedevolution of the mouthparts andor legs for filtering organic material in water(Elouard et al 1990 Gillies et al 1990 Nieto 2003b) A different feeding behaviourlike herbivorous feeding on vascular plants rarely occurs (Gattolliat and Sartori2003)

Convergent evolution can be observed in the different genera living in fast flowIn these genera the median caudal filament is considerably reduced and there arerows of abundant thin setae on the dorsal margin of legs In some cases there is asimple or double carena on the abdominal tergites developed (Waltz and McCafferty1987c Gillies 1991a Gattolliat 2001b) On the contrary other genera likePseudocentroptiloides are considered as psammophilous showing long and slenderlegs and tarsal claws (Glazaczow 1997) The genera Symbiocloeon and Mutelocloeonpresent a unique adaptation among mayflies as their larvae are obligatorycommensals inside mussels (Muller-Liebenau and Heard 1979 Gillies and Elouard1990)

The systematics of Baetidae has been the subject of a lot of attention over the lastthree decades Areas previously poorly known such as Madagascar West AfricaTaiwan or Cuba were subject to intensive sampling and systematic research with thediscovery of several new species and genera During his long life devoted to mayfliesG F Edmunds collected mayflies from almost all over the world His huge collectionwas the source of numerous systematic works led at Purdue University IndianaUSA The genus concept has also evolved greatly firstly by taking into accountlarval characters rather than only imaginal ones and secondly by the use ofphylogenetic methods and the splitting of paraphyletic and polyphyletic genera Theevolution of the global concept of Baetis or Centroptilum and Afroptilum in theAfrotropics perfectly illustrate this new approach (Lugo-Ortiz 1999 Gattolliat et al2008) While the systematic generic classification of Baetidae has been greatlyimproved recently the suprageneric classification of the Baetidae remains unclearHistorically two conflicting concepts have been proposed the division of theBaetidae into different subfamilies (Gillies 1991a) or the gathering of genera inseveral complexes (Waltz et al 1994 Lugo-Ortiz and McCafferty 1996b 1998a1998f) However recent molecular reconstruction showed that the division in

42 J-L Gattolliat and C Nieto

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subfamilies is too simplistic and most of the complexes are not monophyletic(Gattolliat et al 2008)

Recently under the directory of Freshwater Animal Diversity AssessmentProject (FADA) a worldwide global synthesis on Ephemeroptera was publishedproviding the current numbers of known species and generic diversity globallyidentifying the main distribution pattern and biogeographical affinity and high-lighting the main areas of endemicity and possible gaps (Barber-James et al 2008)Subsequently FADA has undertaken to create a worldwide electronic databaseincluding all freshwaters animals For this purpose a database at the specific level ofthe Ephemeroptera was created including the Baetidae (Sartori et al unpublisheddata) Despite this mayfly database being compiled by largely the same authorsthere are some noticeable discrepancies in the number of species per realm especiallyin Palearctic and Nearctic realms While we may expect a few recently describedspecies to be included since publication the main reason for these differences isthe omission of some poorly known species in Barber-James et al (2008)Cross-checking with the electronic database lsquolsquoEphemeroptera of the Worldrsquorsquo ofNikita Kluge (2009) was of great help to fill in this gap for example for species fromthe former USSR or China The results presented in the present study are based onthe Sartori et al database

In the present study we discuss the Baetidae diversity of the differentbiogeographic realms with their rate of endemism and biogeographic affinities Wealso try to show the present state of knowledge of the different faunas and tohighlight those requiring the most attention

Palearctic realm

With 212 species the Palearctic has a high specific diversity (Table 1 Figure 1) Thisdiversity reflects the good knowledge of a great part of the realm rather than a higherdiversity than in other parts of the world The single genus Baetis itself representsabout half of the species part of them having a restricted distribution Most of the 17genera present a wide distribution including other realms The carnivorousRaptobaetopus and the rheophilous Acentrella are the only genera occurring in thePalearctic realm that show important adaptations all other genera are collector-gathers and have no or few adaptations to environmental factors

Four of the eight species of mayflies described by Linnaeus were from thePalearctic realm In the middle of the nineteenth century a few species weredescribed from Western Europe (Rambur 1842 Pictet 1843ndash1845) The contribution

Table 1 Generic and specific diversity by realm Endemicity of taxa endemic from thedifferent realm

No genera Endemicity No species Endemicity

Palearctic 17 12 212 98Afrotropics 40 82 190 100Oriental 23 38 124 99Neotropical 27 66 210 92Neartic 20 25 135 84Australasian 12 50 40 98

Aquatic Insects 43

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of Eaton to the knowledge of European Baetidae was essential in the description ofnew species and a new genus including the various monographs on mayflies hepublished and also with his comprehensive study of the mayflies of England (Eaton1870 Eaton 1871 Eaton 1883ndash1888) His work was outstanding due to the precisionof his drawings and the accuracy of the description He was also the first to have aconcept of the mayfly systematics which is still largely accurate

At the beginning of the twentieth century Bengtsson described several taxa basedon material from Scandinavia (Bengtsson 1912 1914 1917) Some of them werefound to have a much broader distribution in the West Palearctic Mainly based onsingle specimens pinned in different museums Navas described dozens of newspecies from Europe China and Japan (Navas 1911 1913 1917 1931b) Only a fewof them are still considered as valid most of them have never been mentioned sincethe original description Most species described after 1950 from Western Europerepresent alpine or insular vicariants (Thomas and Gazagnes 1984 Thomas 1986Thomas and Soldan 1987 Soldan and Godunko 2006) Several species weredescribed from East Europe (Bogoesco and Tabacaru 1957) and from East Palearctic(Kazlauskas 1963) The fauna of the Himalayas was subject to some attention butmore studies are required (Kapur and Kripalani 1963 Dubey 1971) The Japanesefauna encompasses 11 genera and 39 species most species are endemic to thearchipelago (Fujitani 2008) A great part of them were described before 1950 but themain contribution was made by Gose (1980) with the description of 14 speciesFujitani presented revisions of the Japanese Baetidae and described the larval stagefor several species (Fujitani et al 2003a 2003b) The Baetidae from Korea are alsoquite well-known but mainly included species with a wide distribution (Park et al1996 Bae 1997 Bae and Park 1998) In China several taxa in most of the families ofEphemeroptera were recently described or revised (Zhou et al 2000 Zhou andZheng 2001 Zhou and Braasch 2003 Zhou and Peters 2003 Zhou and Zheng 20032004) but nothing has been undertaken concerning the Baetidae

The study of North African fauna is relatively recent The Moroccan andAlgerian faunas possess several endemic species and show close relationships withWest Palearctic fauna mainly through the Iberian Peninsula (Soldan and Thomas1983a 1983b Thomas and Gagneur 1994 Thomas 1998) Afrotropical affinities arelimited to the single genus Cheleocloeon (Soldan and Thomas 1985) Levant and the

Figure 1 Specific and generic diversity in Palearctic realm Square total number of species(left scale) Triangle total number of genera (right scale)

44 J-L Gattolliat and C Nieto

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Arabian Peninsula have been also subject to attention recently as for North Africathis fauna possesses endemic species but a generic composition mainly similar to theEuropean one (Thomas and Dia 1984 1985 Thomas and Sartori 1989)

Afrotropical realm

The Afrotropical fauna of Baetidae presently encompasses 40 genera and 190 species(Figure 2) they represent about half of the generic and specific diversity of Africanmayflies All of the species and 82 of the genera of Baetidae are endemics from thisrealm the non-endemic genera possess a wide distribution including at least thePalearctic and Oriental realms (Baetis Cloeon and Nigrobaetis for example) TheAfrotropics clearly represent an important centre of diversity for the Baetidae

The distribution of some species covers a great part of the Afrotropics or even inthe case of Cloeon smaeleni the whole realm (Gattolliat and Rabeantoandro 2002)The specific distribution is sometimes greatly underestimated in part due to the lowdegree of knowledge of some faunas but also due to the description of the sametaxon under various names in different areas (Gattolliat unpublished data) Due topeculiar ecological conditions or geographic isolation some taxa have a restricteddistribution Madagascar the Drakensberg and Cape area in South Africa as well asthe volcanic mountain ranges in East and Central Africa are probably the main areasof microendemism

Ulmer (1909) described the first species of Afrotropical Baetidae fromMadagascar and the Comoros Subsequently a few other European researchersdescribed new taxa based on very restricted samples by generalist collectors whichended in European museums (Esben-Petersen 1913 Lestage 1918 Navas 19301931a) Barnard was the first true African ephemeropterist he published the firstmonograph on South African mayflies mainly from Western Cape (Barnard 1932)He was also one of the first who paid attention to the larval stage by rearing mayfliesto obtain larvandashadult association This pioneer study was followed by Crass (1947)and Agnew (1961 1963) contributing to the knowledge of the South African faunaAfter these three studies this fauna already comprised 26 species of Baetidae(compared to the 34 presently known) Most of them were described on a large seriesof specimens from different localities usually both at larval and imaginal stages

Figure 2 Specific and generic diversity in Afrotropical realm Square total number ofspecies (left scale) Triangle total number of genera (right scale)

Aquatic Insects 45

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Kimmins (1956) described a few species from Uganda Demoulin (1964a 1964b1965 1966 1967 1968) reported several species of Baetidae collected duringvarious expeditions In most cases he described them completely but refrained fromnaming them In 1970 he published the first catalogue of the African mayfly faunaincluding all the citations from the Afrotropical area (Demoulin 1970) Based onmore than 20000 imagines Kopelke (1980) described 12 new species from theKalengo mountains near Lake Kivu in Zaire This study was only based on theimaginal stage and the larva-imago association was subsequently possible only for afew species

Gillies contributed greatly to the knowledge of African Baetidae He first workedin East Africa mainly in Tanzania describing several species from this area (Gillies1985 1988 1990 1991a 1991b 1994 1998 Gillies and Wuillot 1997) In the 1970sand 1980s the French ORSTOM team collected aquatic macroinvertebratesrepeatedly in about 100 localities mainly in Guinea Ivory Coast Mali and SenegalThese collections constituted part of an important onchocerciasis controlprogramme (Leveque et al 2003) Elouard and Wuillot in close collaborationwith Gillies investigated this fauna and they described seven genera and about 20species from this area (Gillies 1990 Elouard and Hideux 1991 Wuillot and Gillies1993 1994)

The first mention of Baetidae in Madagascar was provided by Ulmer (1909)Until the middle of the 1990s little work on Malagasy Baetidae had been done atthat time eight species were described but only four of them are now considered asvalid (Gattolliat and Sartori 2003) In less than 10 years 19 genera and 51 specieswere described or reported from this island highlighting the great diversity of thisfauna this faunistic and systematic work was largely based on the material collectedby J-M Elouard and his Malagasy team (Elouard and Gibon 2001) To illustratethe originality and the highly adaptive nature of this fauna we can mention the threeendemic carnivorous genera (Gattolliat and Sartori 2001) the larva of Edmulmeatusgrandis feeding only on vascular plants (Gattolliat and Sartori 2003) and theadaptation for scraping algae from rocks occurring independently in several genera(Lugo-Ortiz and McCafferty 1997 Gattolliat and Sartori 2000 Gattolliat 2001a2001b 2002a 2002b 2004) Compared to other areas of the Afrotropics theMalagasy fauna can be considered as well known even if some systematics problemsneed to be solved and a few new species and genera still await description (Gattolliatand Sartori 2003 Gattolliat et al 2008)

Part of the Arabian Peninsula is also considered as belonging to the Afrotropicalrealm Despite a great part of this area being desert a limited but original faunaoccurs there (Gattolliat and Sartori 2008) This fauna shows direct influence fromboth the Palearctic and Afrotropical realms

In the 1980s and 1990s Waltz and McCafferty then Lugo-Ortiz and McCaffertyrevised the systematics of the Baetidae at the generic level They establishedrespectively three and 14 new genera (Lugo-Ortiz and McCafferty 1999b) It clearlyrepresented an important improvement of our knowledge of Afrotropical Baetidaebut still more systematic work is required before having a complete understanding ofthis fauna In the series lsquolsquoGuides to the Freshwater Invertebrates of SouthernAfricarsquorsquo Barber-James and Lugo-Ortiz (2003) offered a key to the generic level forlarvae of all the Afrotropical mayflies distribution diagnosis and ecology of eachgenus are discussed and an updated version of the checklist is available online(Barber-James 2007)

46 J-L Gattolliat and C Nieto

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We can consider that the faunas of South West and East Africa as well as ofMadagascar are relatively well known In all of these regions there are mostcertainly several species and genera that remain unknown because of restricteddistribution or unusual habitat The fauna of the whole Atlantic side of Africa (fromCameroon to Angola) remains almost completely unknown This fauna probablyencompasses some taxa with a broad distribution like Centroptiloides bifasciata orOphelmatostoma camerunse but it certainly contains several new taxa The area ofthe Ruwenzori mountain range is also poorly known and probably possesses a highdegree of endemism Almost no report of Baetidae is available from NortheasternAfrica north to Kenya

Oriental realm

The first species of Baetidae from the Oriental realm were described from SriLanka and India (Hagen 1858 Eaton 1883ndash1888) (Figure 3) In his work on themayflies of the Sunda Islands Ulmer (1939) provided the first comprehensivereview of an Oriental fauna with complete and accurate descriptions of new speciesat both larval and imaginal stages Kimmins (1947) and Gillies (1949) bothcontributed to the knowledge of the Baetidae from India by respectively describingthree and 11 species from the imaginal stage Apart from these works the Orientalfauna remained very poorly known until the 1980s Muller-Liebenau describedabout 60 new species from the larval stage (about 50 of the species presentlyknown from this area) and six new genera (Muller-Liebenau 1980a 1980b 19811982a 1982b 1982c 1983 1984a 1984b 1985 Muller-Liebenau and Heard 1979Muller-Liebenau and Morihara 1982 Muller-Liebenau and Hubbard 1985) Shemainly described species from Sri Lanka Malaysia and the Philippines Her workconstituted the major contribution to the knowledge of Oriental fauna Waltz andMcCafferty helped to clarify the supra-specific classification by establishing newgenera and complexes of genera (Waltz and McCafferty 1987a 1987c Waltz et al1994)

The Baetidae from Taiwan were the subject of an extensive study allowing thedescription of 14 species (Kang et al 1994 Kang and Yang 1996) The fauna fromrunning waters can be considered as well known but the fauna from still and

Figure 3 Specific and generic diversity in Oriental realm Square total number of species(left scale) Triangle total number of genera (right scale)

Aquatic Insects 47

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standing water remained unknown The fauna of Hong Kong has recently beensubject to close attention with the description of five new species (Tong and Dudgeon2000 2003)

Although some islands like Sri Lanka Taiwan or Hong Kong were the subject ofimportant studies the majority of the Oriental realm remains virtually unknownFor example only provisional checklists of Baetidae at generic level are available forVietnam and Thailand India especially the southern part of this subcontinentprobably possesses a much more diversified fauna than the scattered reports seem toindicate There are also almost no reports from countries with potentially highdiversity such as Cambodia or Myanmar If we consider that in a 85 km2 area inKalimantan (Borneo) intensive sampling allowed the collection of 12 genera ofBaetidae five of them probably being new to science (Sartori et al 2003) we caneasily imagine that only a tiny part of Oriental fauna is presently known Mostfaunas are too poorly known to correctly estimate the distribution of the differentspecimens but preliminary results based on our own collections seem to indicate thatat least some species show a wide distribution including a great part of South EastAsia It is extremely difficult to estimate the specific and generic diversity of theOriental realm even for restricted regions it is probably not overstated to considerthat the species presently known represent less than a half or even a third of the realdiversity

Neotropical realm

With a total surface of 19 million of km2 the Neotropical realm is the fourthbiogeographical region The Baetidae there now encompass 210 species and 27genera (Table 1) The first species of Baetidae from the Neotropics were describedby Pictet (1843ndash1845) Eaton (1871 1883ndash1888) and Weyenbergh (1883)(Figure 4) As for the Afrotropics the descriptions were based on restrictedcollections of imagines deposited in European museums Navas was the mostprolific author unfortunately he described the same species several times underdifferent names he described 26 species of Callibaetis but only eight are presentlyconsidering as valid (Domınguez et al 2006) Most of the species described bythese first authors have been transferred to new genera or synonymised

Figure 4 Specific and generic diversity in Neotropical realm Square total number of species(left scale) Triangle total number of genera (right scale)

48 J-L Gattolliat and C Nieto

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Traver and Edmunds (1968) greatly improved the knowledge of the highlydiversified genus Camelobaetidius Mol (1986) established the only carnivorousgenus for this region Harpagobaetis Lugo-Ortiz and McCafferty (1995 1996a1998c 1999e) described 12 new genera from the Neotropics other generahave been recently established by Nieto (2003b) and McCafferty and Baumgardner(2003) Finally Domınguez et al (2006) published the first comprehensivereview of all South America species of Ephemeroptera with a chapter on thefamily Baetidae including a key illustrations distribution and ecology for eachtaxon

The isolation of this continent for millions of years is one of the mostimportant reasons why 23 of the Neotropical genera and more than 90 of thespecies are endemics from this realm Only seven genera are shared with theNearctic region with a generally higher specific richness in the Neotropics(Apobaetis Baetodes Callibaetis Camelobaetidius and Paracloeodes) only Amer-icabaetis and Baetis are more diversified in the Nearctic (McCafferty 1998) SeveralSouth American species were originally described in the genus Baetis andPseudocloeon but they were all subsequently transferred to other genera such asCamelobaetidius Cloeodes Fallceon and Moribaetis (Lugo-Ortiz and McCafferty1999d McCafferty 2000) Baetis magnus is the only true Baetis which is present inthe Neotropical realm its distribution encompasses a great part of CentralAmerica and the South of USA Baetis is present therefore in all the realmsFallceon certainly also has a Panamerican distribution the generic attribution ofF candidus from Taiwan is clearly a mistake and consequently Fallceon is notpresent in the Oriental realm Cloeodes is in fact the only genus present in SouthAmerica with a wide distribution as species attributed to this genus have beenreported from the whole Pantropical area Most genera possessing an almostworldwide distribution such as Cloeon or LabiobaetisPseudocloeon are onlymissing in the Neotropics this absence also greatly contributes to the originality ofthe Neotropical fauna

Several Neotropical taxa present peculiar adaptations Aturbina is the only genusof the Baetidae without turbinate eyes Callibaetis is one of the few viviparous generain the Ephemeroptera Chane is a spectacular genus with labium and maxillaeextremely well developed for collecting and filtering (Nieto 2003b) presentingsimilarities with the Afroptropical genus Ophelmatostoma (Gillies et al 1990) Mayo(1973) described the first species collected in a waterfall Baetis ellenae latertransferred to Mayobaetis (Waltz and McCafferty 1985) Cloeodes hydation istolerant to short periods of habitat desiccation this species occurs in smalltemporary rock pools (Nolte et al 1996)

Knowledge of the Baetidae from this region has been greatly improved in the lastfew decades new species and genera were described (Dominique et al 2002 Nieto2003a 2003b 2004a 2004b Thomas and Peru 2003) and other genera were reviewed(Nieto 2004c Nieto and Salles 2006 Nieto and Richard 2008) However there arestill regions or even countries with poorly known faunas such as Bolivia Chile PeruEcuador Uruguay and Venezuela these areas almost certainly possess someendemic taxa which remain to be found Recent samplings in Brazilian Amazoniaallowed the discovery of several new species (Salles and Batista 2004 Sallesand Francischetti 2004 Salles et al 2004 2005 Salles 2007 Salles and Polegatto2008) but a great part of this fauna remains unknown (Salles personalcommunication)

Aquatic Insects 49

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Nearctic realm

Being the second largest biogeographic region (229 million km2) the Nearctic regionencompasses only 20 described genera (Table 1) The rate of endemism is also lowOnly five genera have a distribution restricted to this area (25) Seven genera areshared with the Neotropical region other genera present a distributionincluding the Palearctic andor Oriental areas Most genera are collector-gatherersor collector-scrapers no carnivorous Baetidae have been recorded from this regionFrom the 135 Nearctic species 114 are endemic (84) four Holarctic and 17Panamerican

The first ephemeropterologists working in this region were Hagen (1861) Walsh(1863a 1863b) and Eaton (1869 1870 1871 1881 1883ndash1888) Banks (1900 19141918) described eight species of Callibaetis five of which are still valid With 42species McDunnough was the main knowledgeable contributor to Nearctic Baetidae(McDunnough 1922 1923 1924 1925 1926 1929 1931 1932 1936 1939)(Figure 5)

The first accurate revision of all the species from North America was made byNeedham et al (1935) At that time the concept of Baetidae encompassed severaltaxonomic groups that are presently considered as separate families Edmundsrsquo workon mayflies was significant also in this area He proposed one of the first checklists ofmayflies from North America and Mexico (Edmunds and Allen 1957a) Hiscontribution to the biogeography and evolution of Ephemeroptera is still largelyaccurate (Edmunds 1972 1975) In 1976 he published in collaboration with Jensenand Berner a book with all the species known at that time from North and CentralAmerica (Edmunds et al 1976) In the Baetidae he described and reassigned severalspecies (Edmunds 1954 Edmunds and Allen 1957b)

In the Nearctic region contemporary authors such as Jacobus (Jacobus andMcCafferty 2005a 2005b 2006) Lugo-Ortiz and McCafferty (Lugo-Ortiz andMcCafferty 1998d) Waltz (Waltz and McCafferty 1986 1987b 1989) and Wiersema(Wiersema 2000 Wiersema and McCafferty 2004 Wiersema et al 2004) alsocontributed to the advanced knowledge of this fauna by the description of generaand species as well as several revisions with generic reassignments They sustain anactualised web page with a checklist and general information of the mayflies ofNorth and Central America (McCafferty 2009)

Figure 5 Specific and generic diversity in Nearctic realm Square total number of species(left scale) Triangle total number of genera (right scale)

50 J-L Gattolliat and C Nieto

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After 150 years of intensive works the systematics of Baetidae can be consideredas relatively well known even if some taxomomic problems are still not completelysolved The validity of some species needs to be reviewed a few of them have neverbeen mentioned since their original description

Australasian realm

The Australasian realm presently encompasses 40 known species all of themendemic except for Cloeon virens which is also distributed in the Oriental realmTwelve genera occur in this area only six of them being endemic (50) (Table 1)The non-endemic genera are also distributed in the Oriental region The presence ofsome typical Palearctic genera such as Centroptilum in Australasia is doubtful andshould be clarified in the near future

Offadens soror was the first species described from this region by Ulmer in 1908Following Ulmer Tillyard and Harker provided the only significant works onAustralian Baetidae (Tillyard 1936 Harker 1950 1954 1957) and until recently(Figure 6) Suter greatly improved the knowledge of the Ephemeroptera fauna in theAustralian region providing the first key for the larval stage of mayflies (Suter 1979)At that time the Baetidae encompassed only 12 species and five genera andinterestingly Bungona was the only genus which was endemic from this area all theother Australian species being attributed to Palearctic or Holarctic genera Later onSuter described new species and redescribed some genera (Suter 1986 2000 2001Suter and Pearson 2001) He also provided a key specifically for the larvae ofBaetidae which included several undescribed species and genera (Suter 1997)

As in other biogeographic regions Lugo-Ortiz and McCafferty played animportant role by describing several species and four endemic genera from thisregion (Lugo-Ortiz and McCafferty 1998e 1999a 1999c) and by reporting generasuch as Cloeodes or LabiobaetisPseudocloeon (Lugo-Ortiz and McCafferty 1998bLugo-Ortiz et al 1999) They also described several new taxa from New Guinea(Lugo-Ortiz and McCafferty 1999c Lugo-Ortiz et al 1999) Baetidae were one of thetwo families of mayflies reported from the Fiji Islands although it was only apreliminary study the diversity was relatively high with 12 species and three genera(Flowers 1990)

Figure 6 Specific and generic diversity in Australasian realm Square total number ofspecies (left scale) Triangle total number of genera (right scale)

Aquatic Insects 51

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Although the predaceous habit is very unusual in Baetidae two of the 12 generaof this region are carnivorous Echinobaetis from Sulawesi (Mol 1989) andMystaxyops from New Guinea (McCafferty and Sun 2005)

New Zealand is the only important landmass where mayflies are present andBaetidae have never been found This gap remains an enigma These islands are quiteisolated and possess endemic families such as Siphlaenigmatidae and Rallidentidaebut also the widespread family Leptophlebiidae The Baetidae were also consideredto be absent from New Caledonia (Peters 2001) However in material recentlycollected in standing waters half a dozen larvae of Cloeon were included(unpublished data) Because of the absence of Baetidae in previous intensivesamplings and the high power of dispersion of Cloeon it seems likely that a recentcolonisation of the island may be related to human activity If the Leptophlebiidaewere able to remain established on these islands after the break-off of the Gondwanaor to colonise the islands afterwards why were the Baetidae not able to do the sameThe question remains open

Despite recent improvements the fauna of this region remains insufficientlyknown in Australia and even more so on other islands and archipelagos ForAustralia a recent estimation established that only about half of the Australianspecies of Baetidae are described and probably some new genera still need to beestablished (Jeff Webb personal communication)

The biogeographic affinities of this region need to be analysed in detail All thenon-endemic genera are also distributed in the Oriental region they are mainlydiversified in the North East of Australasia and had probably dispersed step by stepthrough South East Asia into Australia where their distribution is limited to theNorth (Lugo-Ortiz and McCafferty 1999b) The genus Cloeodes clearly connects thisarea with the history of Gondwana and the continents placed in the SouthHemisphere Endemic genera such Edmundsiops Offadens and Bungona may berelated to South American taxa (Jeff Webb personal communication)

Conclusion

Since the establishment of the Baetidae in 1815 their knowledge has been hugelyincreased due to the efforts of many authors around the world In 1999 Lugo-Ortizand McCafferty provided the first worldwide synthesis of this family They estimatedthe diversity of this family at 650 species and 87 genera (Lugo-Ortiz and McCafferty1999b) Ten years later the number of species has increased by almost one third toreach 894 species During these 10 years 142 new species were described mainlyfrom South America Madagascar and West Africa It means that the increase of thetotal number of species is mainly due to the description of new species but Lugo-Ortiz and McCafferty had also underestimated the number of species of Baetidae byabout 100 species During these 10 years 17 genera were described and four weresynonymised

Baetidae are mostly diversified and possess a much higher rate of genericendemism in the Tropics and Southern Hemisphere than in the NorthernHemisphere (Barber-James et al 2008) The Afrotropics and Neotropics possessby far the most diversified faunas Despite common Gondwanian origins thesefaunas have almost no affinities The Palearctic fauna shares most genera with theOriental andor Afrotropical faunas as well as the Nearctic fauna Stepwisecolonisation could have occurred through the Palearctic region Central America

52 J-L Gattolliat and C Nieto

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constitutes a bridge between North and South Americas and is under the influence ofboth areas (McCafferty 1998) The affinities of the Nearctic and Neotropical faunasare mainly due to the presence of North American taxa in Central America TheOriental fauna is still only partially known regions of biogeographic importance andwith a high potential diversity such as the Indian subcontinent and South East Asiaremain almost completely unknown Local studies clearly indicate that the diversityis quite high but the distribution of the taxa remains unknown The Oriental realm istherefore probably the most promising and challenging area Even if the globaldiversity is probably lower in Australasia the lack of knowledge of the fauna isobvious especially in the tropical part of this area such as New Guinea

The knowledge on systematics and faunistic composition are the first inevitablesteps to understand and preserve a natural system Most of the remaining unknownor poorly known faunas are under a direct human threat This underlines theurgency of this work before these faunas are definitively lost this is especially true inmost of the tropical rain forest areas where a primordial aquatic fauna is still presentFor areas with sufficient knowledge it is also the duty of the systematists to offer thenecessary tools for a correct identification of the taxa We can only encourageeveryone to make our knowledge accessible to others scientists especially by creatingkeys and checklists at different systematic levels

A comprehensive reconstruction of the phylogeny of the Baetidae is also highlynecessary Such a reconstruction should be based on modern tools and include bothmolecular and morphological characters It is the only way to solve contradictorysuprageneric relationships Such studies would allow a more accurate understandingof the relationships between the different areas in a global biogeographic approachThey would help to understand the historical influence of Gondwana and Laurasiaon the mayfly fauna

Acknowledgements

We want to express our appreciation to Michel Sartori (Museum of Zoology LausanneSwitzerland) and Helen Barber-James (Albany Museum Grahamstown South Africa) foruseful comments on this paper and great help in the conception of the Baetidae databaseAline Pasche (Museum of Zoology Lausanne Switzerland) offered technical supportFinancial support from the Argentine National Council of Scientific and TechnologicalResearch (CONICET) and ANPCYT (Pict 524-Pict 528) to which the junior author belongsare acknowledged The electronic database lsquolsquoEphemeroptera of the Worldrsquorsquo of Nikita Kluge(httpwwwinsectabiopuruzEph-sppContentshtm) was of great help for cross-checkingof our own database

References

Agnew JD (1961) lsquoNew Baetidae (Ephem) from South Africarsquo Novos Taxa Entomologicos25 3ndash18

Agnew JD (1963) lsquoNew South African records of imperfectly known Baetidae (Ephem)rsquoHydrobiologia 22 41ndash46

Bae YJ (1997) lsquoTaxonomy of Cloeon and Procloeon (Ephemeroptera Baetidae) in KorearsquoThe Korean Journal of Systematics Biology 13 303ndash314

Bae YJ and Park SY (1998) lsquoAlainites Baetis Labiobaetis and Nigrobaetis(Ephemeroptera Baetidae) in Korearsquo The Korean Journal of Systematics Biology 141ndash12

Banks N (1900) lsquoNew genera and species of Nearctic Neuropteroid Insectsrsquo Transactions ofthe American Entomological Society 26 239ndash259

Banks N (1914) lsquoNew neuropteroid insects native and exoticrsquo Proceedings of the Academy ofNatural Sciences of Philadelphia 66 608ndash632

Aquatic Insects 53

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reduced or absent three-segmented mid and hind tarsi and membranous penis(Edmunds et al 1976) Larvae are also unique among mayflies in having the lateralbranches of the epicranial suture anterior to the lateral ocelli and the ventralorientation of the dorsal lobe at the apex of femora (Wang and McCafferty 1996)Larvae are pisciform they generally possess long antennae and simple or doubleovoid gills on segments IndashVII or IIndashVII

Baetidae are very common in any kind of freshwater they are mainly diversifiedin unpolluted running water especially in the tropics Although they are lessdiversified in standing waters with genera like Cloeon the Baetidae constitute animportant part of the insect biomass in ponds Most species of Baetidae arecollector-gatherers feeding mainly on detritus Unusual adaptations to peculiar dietsoccur in a few genera Twelve species belonging to nine genera are considered to becarnivorous They feed mainly on larvae of Diptera but some are specialised infeeding on other mayflies (Gattolliat and Sartori 2001 McCafferty and Sun 2005)Although these carnivorous genera present similar adaptations of the mouthpartsthey clearly belong to different lineages (Monaghan et al 2005) In several lineagesspecies or genera show adaptations for scrapping algae on the top of stones (Lugo-Ortiz and McCafferty 1997 Gattolliat 2001b) Some genera have an advancedevolution of the mouthparts andor legs for filtering organic material in water(Elouard et al 1990 Gillies et al 1990 Nieto 2003b) A different feeding behaviourlike herbivorous feeding on vascular plants rarely occurs (Gattolliat and Sartori2003)

Convergent evolution can be observed in the different genera living in fast flowIn these genera the median caudal filament is considerably reduced and there arerows of abundant thin setae on the dorsal margin of legs In some cases there is asimple or double carena on the abdominal tergites developed (Waltz and McCafferty1987c Gillies 1991a Gattolliat 2001b) On the contrary other genera likePseudocentroptiloides are considered as psammophilous showing long and slenderlegs and tarsal claws (Glazaczow 1997) The genera Symbiocloeon and Mutelocloeonpresent a unique adaptation among mayflies as their larvae are obligatorycommensals inside mussels (Muller-Liebenau and Heard 1979 Gillies and Elouard1990)

The systematics of Baetidae has been the subject of a lot of attention over the lastthree decades Areas previously poorly known such as Madagascar West AfricaTaiwan or Cuba were subject to intensive sampling and systematic research with thediscovery of several new species and genera During his long life devoted to mayfliesG F Edmunds collected mayflies from almost all over the world His huge collectionwas the source of numerous systematic works led at Purdue University IndianaUSA The genus concept has also evolved greatly firstly by taking into accountlarval characters rather than only imaginal ones and secondly by the use ofphylogenetic methods and the splitting of paraphyletic and polyphyletic genera Theevolution of the global concept of Baetis or Centroptilum and Afroptilum in theAfrotropics perfectly illustrate this new approach (Lugo-Ortiz 1999 Gattolliat et al2008) While the systematic generic classification of Baetidae has been greatlyimproved recently the suprageneric classification of the Baetidae remains unclearHistorically two conflicting concepts have been proposed the division of theBaetidae into different subfamilies (Gillies 1991a) or the gathering of genera inseveral complexes (Waltz et al 1994 Lugo-Ortiz and McCafferty 1996b 1998a1998f) However recent molecular reconstruction showed that the division in

42 J-L Gattolliat and C Nieto

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subfamilies is too simplistic and most of the complexes are not monophyletic(Gattolliat et al 2008)

Recently under the directory of Freshwater Animal Diversity AssessmentProject (FADA) a worldwide global synthesis on Ephemeroptera was publishedproviding the current numbers of known species and generic diversity globallyidentifying the main distribution pattern and biogeographical affinity and high-lighting the main areas of endemicity and possible gaps (Barber-James et al 2008)Subsequently FADA has undertaken to create a worldwide electronic databaseincluding all freshwaters animals For this purpose a database at the specific level ofthe Ephemeroptera was created including the Baetidae (Sartori et al unpublisheddata) Despite this mayfly database being compiled by largely the same authorsthere are some noticeable discrepancies in the number of species per realm especiallyin Palearctic and Nearctic realms While we may expect a few recently describedspecies to be included since publication the main reason for these differences isthe omission of some poorly known species in Barber-James et al (2008)Cross-checking with the electronic database lsquolsquoEphemeroptera of the Worldrsquorsquo ofNikita Kluge (2009) was of great help to fill in this gap for example for species fromthe former USSR or China The results presented in the present study are based onthe Sartori et al database

In the present study we discuss the Baetidae diversity of the differentbiogeographic realms with their rate of endemism and biogeographic affinities Wealso try to show the present state of knowledge of the different faunas and tohighlight those requiring the most attention

Palearctic realm

With 212 species the Palearctic has a high specific diversity (Table 1 Figure 1) Thisdiversity reflects the good knowledge of a great part of the realm rather than a higherdiversity than in other parts of the world The single genus Baetis itself representsabout half of the species part of them having a restricted distribution Most of the 17genera present a wide distribution including other realms The carnivorousRaptobaetopus and the rheophilous Acentrella are the only genera occurring in thePalearctic realm that show important adaptations all other genera are collector-gathers and have no or few adaptations to environmental factors

Four of the eight species of mayflies described by Linnaeus were from thePalearctic realm In the middle of the nineteenth century a few species weredescribed from Western Europe (Rambur 1842 Pictet 1843ndash1845) The contribution

Table 1 Generic and specific diversity by realm Endemicity of taxa endemic from thedifferent realm

No genera Endemicity No species Endemicity

Palearctic 17 12 212 98Afrotropics 40 82 190 100Oriental 23 38 124 99Neotropical 27 66 210 92Neartic 20 25 135 84Australasian 12 50 40 98

Aquatic Insects 43

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of Eaton to the knowledge of European Baetidae was essential in the description ofnew species and a new genus including the various monographs on mayflies hepublished and also with his comprehensive study of the mayflies of England (Eaton1870 Eaton 1871 Eaton 1883ndash1888) His work was outstanding due to the precisionof his drawings and the accuracy of the description He was also the first to have aconcept of the mayfly systematics which is still largely accurate

At the beginning of the twentieth century Bengtsson described several taxa basedon material from Scandinavia (Bengtsson 1912 1914 1917) Some of them werefound to have a much broader distribution in the West Palearctic Mainly based onsingle specimens pinned in different museums Navas described dozens of newspecies from Europe China and Japan (Navas 1911 1913 1917 1931b) Only a fewof them are still considered as valid most of them have never been mentioned sincethe original description Most species described after 1950 from Western Europerepresent alpine or insular vicariants (Thomas and Gazagnes 1984 Thomas 1986Thomas and Soldan 1987 Soldan and Godunko 2006) Several species weredescribed from East Europe (Bogoesco and Tabacaru 1957) and from East Palearctic(Kazlauskas 1963) The fauna of the Himalayas was subject to some attention butmore studies are required (Kapur and Kripalani 1963 Dubey 1971) The Japanesefauna encompasses 11 genera and 39 species most species are endemic to thearchipelago (Fujitani 2008) A great part of them were described before 1950 but themain contribution was made by Gose (1980) with the description of 14 speciesFujitani presented revisions of the Japanese Baetidae and described the larval stagefor several species (Fujitani et al 2003a 2003b) The Baetidae from Korea are alsoquite well-known but mainly included species with a wide distribution (Park et al1996 Bae 1997 Bae and Park 1998) In China several taxa in most of the families ofEphemeroptera were recently described or revised (Zhou et al 2000 Zhou andZheng 2001 Zhou and Braasch 2003 Zhou and Peters 2003 Zhou and Zheng 20032004) but nothing has been undertaken concerning the Baetidae

The study of North African fauna is relatively recent The Moroccan andAlgerian faunas possess several endemic species and show close relationships withWest Palearctic fauna mainly through the Iberian Peninsula (Soldan and Thomas1983a 1983b Thomas and Gagneur 1994 Thomas 1998) Afrotropical affinities arelimited to the single genus Cheleocloeon (Soldan and Thomas 1985) Levant and the

Figure 1 Specific and generic diversity in Palearctic realm Square total number of species(left scale) Triangle total number of genera (right scale)

44 J-L Gattolliat and C Nieto

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Arabian Peninsula have been also subject to attention recently as for North Africathis fauna possesses endemic species but a generic composition mainly similar to theEuropean one (Thomas and Dia 1984 1985 Thomas and Sartori 1989)

Afrotropical realm

The Afrotropical fauna of Baetidae presently encompasses 40 genera and 190 species(Figure 2) they represent about half of the generic and specific diversity of Africanmayflies All of the species and 82 of the genera of Baetidae are endemics from thisrealm the non-endemic genera possess a wide distribution including at least thePalearctic and Oriental realms (Baetis Cloeon and Nigrobaetis for example) TheAfrotropics clearly represent an important centre of diversity for the Baetidae

The distribution of some species covers a great part of the Afrotropics or even inthe case of Cloeon smaeleni the whole realm (Gattolliat and Rabeantoandro 2002)The specific distribution is sometimes greatly underestimated in part due to the lowdegree of knowledge of some faunas but also due to the description of the sametaxon under various names in different areas (Gattolliat unpublished data) Due topeculiar ecological conditions or geographic isolation some taxa have a restricteddistribution Madagascar the Drakensberg and Cape area in South Africa as well asthe volcanic mountain ranges in East and Central Africa are probably the main areasof microendemism

Ulmer (1909) described the first species of Afrotropical Baetidae fromMadagascar and the Comoros Subsequently a few other European researchersdescribed new taxa based on very restricted samples by generalist collectors whichended in European museums (Esben-Petersen 1913 Lestage 1918 Navas 19301931a) Barnard was the first true African ephemeropterist he published the firstmonograph on South African mayflies mainly from Western Cape (Barnard 1932)He was also one of the first who paid attention to the larval stage by rearing mayfliesto obtain larvandashadult association This pioneer study was followed by Crass (1947)and Agnew (1961 1963) contributing to the knowledge of the South African faunaAfter these three studies this fauna already comprised 26 species of Baetidae(compared to the 34 presently known) Most of them were described on a large seriesof specimens from different localities usually both at larval and imaginal stages

Figure 2 Specific and generic diversity in Afrotropical realm Square total number ofspecies (left scale) Triangle total number of genera (right scale)

Aquatic Insects 45

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Kimmins (1956) described a few species from Uganda Demoulin (1964a 1964b1965 1966 1967 1968) reported several species of Baetidae collected duringvarious expeditions In most cases he described them completely but refrained fromnaming them In 1970 he published the first catalogue of the African mayfly faunaincluding all the citations from the Afrotropical area (Demoulin 1970) Based onmore than 20000 imagines Kopelke (1980) described 12 new species from theKalengo mountains near Lake Kivu in Zaire This study was only based on theimaginal stage and the larva-imago association was subsequently possible only for afew species

Gillies contributed greatly to the knowledge of African Baetidae He first workedin East Africa mainly in Tanzania describing several species from this area (Gillies1985 1988 1990 1991a 1991b 1994 1998 Gillies and Wuillot 1997) In the 1970sand 1980s the French ORSTOM team collected aquatic macroinvertebratesrepeatedly in about 100 localities mainly in Guinea Ivory Coast Mali and SenegalThese collections constituted part of an important onchocerciasis controlprogramme (Leveque et al 2003) Elouard and Wuillot in close collaborationwith Gillies investigated this fauna and they described seven genera and about 20species from this area (Gillies 1990 Elouard and Hideux 1991 Wuillot and Gillies1993 1994)

The first mention of Baetidae in Madagascar was provided by Ulmer (1909)Until the middle of the 1990s little work on Malagasy Baetidae had been done atthat time eight species were described but only four of them are now considered asvalid (Gattolliat and Sartori 2003) In less than 10 years 19 genera and 51 specieswere described or reported from this island highlighting the great diversity of thisfauna this faunistic and systematic work was largely based on the material collectedby J-M Elouard and his Malagasy team (Elouard and Gibon 2001) To illustratethe originality and the highly adaptive nature of this fauna we can mention the threeendemic carnivorous genera (Gattolliat and Sartori 2001) the larva of Edmulmeatusgrandis feeding only on vascular plants (Gattolliat and Sartori 2003) and theadaptation for scraping algae from rocks occurring independently in several genera(Lugo-Ortiz and McCafferty 1997 Gattolliat and Sartori 2000 Gattolliat 2001a2001b 2002a 2002b 2004) Compared to other areas of the Afrotropics theMalagasy fauna can be considered as well known even if some systematics problemsneed to be solved and a few new species and genera still await description (Gattolliatand Sartori 2003 Gattolliat et al 2008)

Part of the Arabian Peninsula is also considered as belonging to the Afrotropicalrealm Despite a great part of this area being desert a limited but original faunaoccurs there (Gattolliat and Sartori 2008) This fauna shows direct influence fromboth the Palearctic and Afrotropical realms

In the 1980s and 1990s Waltz and McCafferty then Lugo-Ortiz and McCaffertyrevised the systematics of the Baetidae at the generic level They establishedrespectively three and 14 new genera (Lugo-Ortiz and McCafferty 1999b) It clearlyrepresented an important improvement of our knowledge of Afrotropical Baetidaebut still more systematic work is required before having a complete understanding ofthis fauna In the series lsquolsquoGuides to the Freshwater Invertebrates of SouthernAfricarsquorsquo Barber-James and Lugo-Ortiz (2003) offered a key to the generic level forlarvae of all the Afrotropical mayflies distribution diagnosis and ecology of eachgenus are discussed and an updated version of the checklist is available online(Barber-James 2007)

46 J-L Gattolliat and C Nieto

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We can consider that the faunas of South West and East Africa as well as ofMadagascar are relatively well known In all of these regions there are mostcertainly several species and genera that remain unknown because of restricteddistribution or unusual habitat The fauna of the whole Atlantic side of Africa (fromCameroon to Angola) remains almost completely unknown This fauna probablyencompasses some taxa with a broad distribution like Centroptiloides bifasciata orOphelmatostoma camerunse but it certainly contains several new taxa The area ofthe Ruwenzori mountain range is also poorly known and probably possesses a highdegree of endemism Almost no report of Baetidae is available from NortheasternAfrica north to Kenya

Oriental realm

The first species of Baetidae from the Oriental realm were described from SriLanka and India (Hagen 1858 Eaton 1883ndash1888) (Figure 3) In his work on themayflies of the Sunda Islands Ulmer (1939) provided the first comprehensivereview of an Oriental fauna with complete and accurate descriptions of new speciesat both larval and imaginal stages Kimmins (1947) and Gillies (1949) bothcontributed to the knowledge of the Baetidae from India by respectively describingthree and 11 species from the imaginal stage Apart from these works the Orientalfauna remained very poorly known until the 1980s Muller-Liebenau describedabout 60 new species from the larval stage (about 50 of the species presentlyknown from this area) and six new genera (Muller-Liebenau 1980a 1980b 19811982a 1982b 1982c 1983 1984a 1984b 1985 Muller-Liebenau and Heard 1979Muller-Liebenau and Morihara 1982 Muller-Liebenau and Hubbard 1985) Shemainly described species from Sri Lanka Malaysia and the Philippines Her workconstituted the major contribution to the knowledge of Oriental fauna Waltz andMcCafferty helped to clarify the supra-specific classification by establishing newgenera and complexes of genera (Waltz and McCafferty 1987a 1987c Waltz et al1994)

The Baetidae from Taiwan were the subject of an extensive study allowing thedescription of 14 species (Kang et al 1994 Kang and Yang 1996) The fauna fromrunning waters can be considered as well known but the fauna from still and

Figure 3 Specific and generic diversity in Oriental realm Square total number of species(left scale) Triangle total number of genera (right scale)

Aquatic Insects 47

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standing water remained unknown The fauna of Hong Kong has recently beensubject to close attention with the description of five new species (Tong and Dudgeon2000 2003)

Although some islands like Sri Lanka Taiwan or Hong Kong were the subject ofimportant studies the majority of the Oriental realm remains virtually unknownFor example only provisional checklists of Baetidae at generic level are available forVietnam and Thailand India especially the southern part of this subcontinentprobably possesses a much more diversified fauna than the scattered reports seem toindicate There are also almost no reports from countries with potentially highdiversity such as Cambodia or Myanmar If we consider that in a 85 km2 area inKalimantan (Borneo) intensive sampling allowed the collection of 12 genera ofBaetidae five of them probably being new to science (Sartori et al 2003) we caneasily imagine that only a tiny part of Oriental fauna is presently known Mostfaunas are too poorly known to correctly estimate the distribution of the differentspecimens but preliminary results based on our own collections seem to indicate thatat least some species show a wide distribution including a great part of South EastAsia It is extremely difficult to estimate the specific and generic diversity of theOriental realm even for restricted regions it is probably not overstated to considerthat the species presently known represent less than a half or even a third of the realdiversity

Neotropical realm

With a total surface of 19 million of km2 the Neotropical realm is the fourthbiogeographical region The Baetidae there now encompass 210 species and 27genera (Table 1) The first species of Baetidae from the Neotropics were describedby Pictet (1843ndash1845) Eaton (1871 1883ndash1888) and Weyenbergh (1883)(Figure 4) As for the Afrotropics the descriptions were based on restrictedcollections of imagines deposited in European museums Navas was the mostprolific author unfortunately he described the same species several times underdifferent names he described 26 species of Callibaetis but only eight are presentlyconsidering as valid (Domınguez et al 2006) Most of the species described bythese first authors have been transferred to new genera or synonymised

Figure 4 Specific and generic diversity in Neotropical realm Square total number of species(left scale) Triangle total number of genera (right scale)

48 J-L Gattolliat and C Nieto

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Traver and Edmunds (1968) greatly improved the knowledge of the highlydiversified genus Camelobaetidius Mol (1986) established the only carnivorousgenus for this region Harpagobaetis Lugo-Ortiz and McCafferty (1995 1996a1998c 1999e) described 12 new genera from the Neotropics other generahave been recently established by Nieto (2003b) and McCafferty and Baumgardner(2003) Finally Domınguez et al (2006) published the first comprehensivereview of all South America species of Ephemeroptera with a chapter on thefamily Baetidae including a key illustrations distribution and ecology for eachtaxon

The isolation of this continent for millions of years is one of the mostimportant reasons why 23 of the Neotropical genera and more than 90 of thespecies are endemics from this realm Only seven genera are shared with theNearctic region with a generally higher specific richness in the Neotropics(Apobaetis Baetodes Callibaetis Camelobaetidius and Paracloeodes) only Amer-icabaetis and Baetis are more diversified in the Nearctic (McCafferty 1998) SeveralSouth American species were originally described in the genus Baetis andPseudocloeon but they were all subsequently transferred to other genera such asCamelobaetidius Cloeodes Fallceon and Moribaetis (Lugo-Ortiz and McCafferty1999d McCafferty 2000) Baetis magnus is the only true Baetis which is present inthe Neotropical realm its distribution encompasses a great part of CentralAmerica and the South of USA Baetis is present therefore in all the realmsFallceon certainly also has a Panamerican distribution the generic attribution ofF candidus from Taiwan is clearly a mistake and consequently Fallceon is notpresent in the Oriental realm Cloeodes is in fact the only genus present in SouthAmerica with a wide distribution as species attributed to this genus have beenreported from the whole Pantropical area Most genera possessing an almostworldwide distribution such as Cloeon or LabiobaetisPseudocloeon are onlymissing in the Neotropics this absence also greatly contributes to the originality ofthe Neotropical fauna

Several Neotropical taxa present peculiar adaptations Aturbina is the only genusof the Baetidae without turbinate eyes Callibaetis is one of the few viviparous generain the Ephemeroptera Chane is a spectacular genus with labium and maxillaeextremely well developed for collecting and filtering (Nieto 2003b) presentingsimilarities with the Afroptropical genus Ophelmatostoma (Gillies et al 1990) Mayo(1973) described the first species collected in a waterfall Baetis ellenae latertransferred to Mayobaetis (Waltz and McCafferty 1985) Cloeodes hydation istolerant to short periods of habitat desiccation this species occurs in smalltemporary rock pools (Nolte et al 1996)

Knowledge of the Baetidae from this region has been greatly improved in the lastfew decades new species and genera were described (Dominique et al 2002 Nieto2003a 2003b 2004a 2004b Thomas and Peru 2003) and other genera were reviewed(Nieto 2004c Nieto and Salles 2006 Nieto and Richard 2008) However there arestill regions or even countries with poorly known faunas such as Bolivia Chile PeruEcuador Uruguay and Venezuela these areas almost certainly possess someendemic taxa which remain to be found Recent samplings in Brazilian Amazoniaallowed the discovery of several new species (Salles and Batista 2004 Sallesand Francischetti 2004 Salles et al 2004 2005 Salles 2007 Salles and Polegatto2008) but a great part of this fauna remains unknown (Salles personalcommunication)

Aquatic Insects 49

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Nearctic realm

Being the second largest biogeographic region (229 million km2) the Nearctic regionencompasses only 20 described genera (Table 1) The rate of endemism is also lowOnly five genera have a distribution restricted to this area (25) Seven genera areshared with the Neotropical region other genera present a distributionincluding the Palearctic andor Oriental areas Most genera are collector-gatherersor collector-scrapers no carnivorous Baetidae have been recorded from this regionFrom the 135 Nearctic species 114 are endemic (84) four Holarctic and 17Panamerican

The first ephemeropterologists working in this region were Hagen (1861) Walsh(1863a 1863b) and Eaton (1869 1870 1871 1881 1883ndash1888) Banks (1900 19141918) described eight species of Callibaetis five of which are still valid With 42species McDunnough was the main knowledgeable contributor to Nearctic Baetidae(McDunnough 1922 1923 1924 1925 1926 1929 1931 1932 1936 1939)(Figure 5)

The first accurate revision of all the species from North America was made byNeedham et al (1935) At that time the concept of Baetidae encompassed severaltaxonomic groups that are presently considered as separate families Edmundsrsquo workon mayflies was significant also in this area He proposed one of the first checklists ofmayflies from North America and Mexico (Edmunds and Allen 1957a) Hiscontribution to the biogeography and evolution of Ephemeroptera is still largelyaccurate (Edmunds 1972 1975) In 1976 he published in collaboration with Jensenand Berner a book with all the species known at that time from North and CentralAmerica (Edmunds et al 1976) In the Baetidae he described and reassigned severalspecies (Edmunds 1954 Edmunds and Allen 1957b)

In the Nearctic region contemporary authors such as Jacobus (Jacobus andMcCafferty 2005a 2005b 2006) Lugo-Ortiz and McCafferty (Lugo-Ortiz andMcCafferty 1998d) Waltz (Waltz and McCafferty 1986 1987b 1989) and Wiersema(Wiersema 2000 Wiersema and McCafferty 2004 Wiersema et al 2004) alsocontributed to the advanced knowledge of this fauna by the description of generaand species as well as several revisions with generic reassignments They sustain anactualised web page with a checklist and general information of the mayflies ofNorth and Central America (McCafferty 2009)

Figure 5 Specific and generic diversity in Nearctic realm Square total number of species(left scale) Triangle total number of genera (right scale)

50 J-L Gattolliat and C Nieto

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After 150 years of intensive works the systematics of Baetidae can be consideredas relatively well known even if some taxomomic problems are still not completelysolved The validity of some species needs to be reviewed a few of them have neverbeen mentioned since their original description

Australasian realm

The Australasian realm presently encompasses 40 known species all of themendemic except for Cloeon virens which is also distributed in the Oriental realmTwelve genera occur in this area only six of them being endemic (50) (Table 1)The non-endemic genera are also distributed in the Oriental region The presence ofsome typical Palearctic genera such as Centroptilum in Australasia is doubtful andshould be clarified in the near future

Offadens soror was the first species described from this region by Ulmer in 1908Following Ulmer Tillyard and Harker provided the only significant works onAustralian Baetidae (Tillyard 1936 Harker 1950 1954 1957) and until recently(Figure 6) Suter greatly improved the knowledge of the Ephemeroptera fauna in theAustralian region providing the first key for the larval stage of mayflies (Suter 1979)At that time the Baetidae encompassed only 12 species and five genera andinterestingly Bungona was the only genus which was endemic from this area all theother Australian species being attributed to Palearctic or Holarctic genera Later onSuter described new species and redescribed some genera (Suter 1986 2000 2001Suter and Pearson 2001) He also provided a key specifically for the larvae ofBaetidae which included several undescribed species and genera (Suter 1997)

As in other biogeographic regions Lugo-Ortiz and McCafferty played animportant role by describing several species and four endemic genera from thisregion (Lugo-Ortiz and McCafferty 1998e 1999a 1999c) and by reporting generasuch as Cloeodes or LabiobaetisPseudocloeon (Lugo-Ortiz and McCafferty 1998bLugo-Ortiz et al 1999) They also described several new taxa from New Guinea(Lugo-Ortiz and McCafferty 1999c Lugo-Ortiz et al 1999) Baetidae were one of thetwo families of mayflies reported from the Fiji Islands although it was only apreliminary study the diversity was relatively high with 12 species and three genera(Flowers 1990)

Figure 6 Specific and generic diversity in Australasian realm Square total number ofspecies (left scale) Triangle total number of genera (right scale)

Aquatic Insects 51

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Although the predaceous habit is very unusual in Baetidae two of the 12 generaof this region are carnivorous Echinobaetis from Sulawesi (Mol 1989) andMystaxyops from New Guinea (McCafferty and Sun 2005)

New Zealand is the only important landmass where mayflies are present andBaetidae have never been found This gap remains an enigma These islands are quiteisolated and possess endemic families such as Siphlaenigmatidae and Rallidentidaebut also the widespread family Leptophlebiidae The Baetidae were also consideredto be absent from New Caledonia (Peters 2001) However in material recentlycollected in standing waters half a dozen larvae of Cloeon were included(unpublished data) Because of the absence of Baetidae in previous intensivesamplings and the high power of dispersion of Cloeon it seems likely that a recentcolonisation of the island may be related to human activity If the Leptophlebiidaewere able to remain established on these islands after the break-off of the Gondwanaor to colonise the islands afterwards why were the Baetidae not able to do the sameThe question remains open

Despite recent improvements the fauna of this region remains insufficientlyknown in Australia and even more so on other islands and archipelagos ForAustralia a recent estimation established that only about half of the Australianspecies of Baetidae are described and probably some new genera still need to beestablished (Jeff Webb personal communication)

The biogeographic affinities of this region need to be analysed in detail All thenon-endemic genera are also distributed in the Oriental region they are mainlydiversified in the North East of Australasia and had probably dispersed step by stepthrough South East Asia into Australia where their distribution is limited to theNorth (Lugo-Ortiz and McCafferty 1999b) The genus Cloeodes clearly connects thisarea with the history of Gondwana and the continents placed in the SouthHemisphere Endemic genera such Edmundsiops Offadens and Bungona may berelated to South American taxa (Jeff Webb personal communication)

Conclusion

Since the establishment of the Baetidae in 1815 their knowledge has been hugelyincreased due to the efforts of many authors around the world In 1999 Lugo-Ortizand McCafferty provided the first worldwide synthesis of this family They estimatedthe diversity of this family at 650 species and 87 genera (Lugo-Ortiz and McCafferty1999b) Ten years later the number of species has increased by almost one third toreach 894 species During these 10 years 142 new species were described mainlyfrom South America Madagascar and West Africa It means that the increase of thetotal number of species is mainly due to the description of new species but Lugo-Ortiz and McCafferty had also underestimated the number of species of Baetidae byabout 100 species During these 10 years 17 genera were described and four weresynonymised

Baetidae are mostly diversified and possess a much higher rate of genericendemism in the Tropics and Southern Hemisphere than in the NorthernHemisphere (Barber-James et al 2008) The Afrotropics and Neotropics possessby far the most diversified faunas Despite common Gondwanian origins thesefaunas have almost no affinities The Palearctic fauna shares most genera with theOriental andor Afrotropical faunas as well as the Nearctic fauna Stepwisecolonisation could have occurred through the Palearctic region Central America

52 J-L Gattolliat and C Nieto

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constitutes a bridge between North and South Americas and is under the influence ofboth areas (McCafferty 1998) The affinities of the Nearctic and Neotropical faunasare mainly due to the presence of North American taxa in Central America TheOriental fauna is still only partially known regions of biogeographic importance andwith a high potential diversity such as the Indian subcontinent and South East Asiaremain almost completely unknown Local studies clearly indicate that the diversityis quite high but the distribution of the taxa remains unknown The Oriental realm istherefore probably the most promising and challenging area Even if the globaldiversity is probably lower in Australasia the lack of knowledge of the fauna isobvious especially in the tropical part of this area such as New Guinea

The knowledge on systematics and faunistic composition are the first inevitablesteps to understand and preserve a natural system Most of the remaining unknownor poorly known faunas are under a direct human threat This underlines theurgency of this work before these faunas are definitively lost this is especially true inmost of the tropical rain forest areas where a primordial aquatic fauna is still presentFor areas with sufficient knowledge it is also the duty of the systematists to offer thenecessary tools for a correct identification of the taxa We can only encourageeveryone to make our knowledge accessible to others scientists especially by creatingkeys and checklists at different systematic levels

A comprehensive reconstruction of the phylogeny of the Baetidae is also highlynecessary Such a reconstruction should be based on modern tools and include bothmolecular and morphological characters It is the only way to solve contradictorysuprageneric relationships Such studies would allow a more accurate understandingof the relationships between the different areas in a global biogeographic approachThey would help to understand the historical influence of Gondwana and Laurasiaon the mayfly fauna

Acknowledgements

We want to express our appreciation to Michel Sartori (Museum of Zoology LausanneSwitzerland) and Helen Barber-James (Albany Museum Grahamstown South Africa) foruseful comments on this paper and great help in the conception of the Baetidae databaseAline Pasche (Museum of Zoology Lausanne Switzerland) offered technical supportFinancial support from the Argentine National Council of Scientific and TechnologicalResearch (CONICET) and ANPCYT (Pict 524-Pict 528) to which the junior author belongsare acknowledged The electronic database lsquolsquoEphemeroptera of the Worldrsquorsquo of Nikita Kluge(httpwwwinsectabiopuruzEph-sppContentshtm) was of great help for cross-checkingof our own database

References

Agnew JD (1961) lsquoNew Baetidae (Ephem) from South Africarsquo Novos Taxa Entomologicos25 3ndash18

Agnew JD (1963) lsquoNew South African records of imperfectly known Baetidae (Ephem)rsquoHydrobiologia 22 41ndash46

Bae YJ (1997) lsquoTaxonomy of Cloeon and Procloeon (Ephemeroptera Baetidae) in KorearsquoThe Korean Journal of Systematics Biology 13 303ndash314

Bae YJ and Park SY (1998) lsquoAlainites Baetis Labiobaetis and Nigrobaetis(Ephemeroptera Baetidae) in Korearsquo The Korean Journal of Systematics Biology 141ndash12

Banks N (1900) lsquoNew genera and species of Nearctic Neuropteroid Insectsrsquo Transactions ofthe American Entomological Society 26 239ndash259

Banks N (1914) lsquoNew neuropteroid insects native and exoticrsquo Proceedings of the Academy ofNatural Sciences of Philadelphia 66 608ndash632

Aquatic Insects 53

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subfamilies is too simplistic and most of the complexes are not monophyletic(Gattolliat et al 2008)

Recently under the directory of Freshwater Animal Diversity AssessmentProject (FADA) a worldwide global synthesis on Ephemeroptera was publishedproviding the current numbers of known species and generic diversity globallyidentifying the main distribution pattern and biogeographical affinity and high-lighting the main areas of endemicity and possible gaps (Barber-James et al 2008)Subsequently FADA has undertaken to create a worldwide electronic databaseincluding all freshwaters animals For this purpose a database at the specific level ofthe Ephemeroptera was created including the Baetidae (Sartori et al unpublisheddata) Despite this mayfly database being compiled by largely the same authorsthere are some noticeable discrepancies in the number of species per realm especiallyin Palearctic and Nearctic realms While we may expect a few recently describedspecies to be included since publication the main reason for these differences isthe omission of some poorly known species in Barber-James et al (2008)Cross-checking with the electronic database lsquolsquoEphemeroptera of the Worldrsquorsquo ofNikita Kluge (2009) was of great help to fill in this gap for example for species fromthe former USSR or China The results presented in the present study are based onthe Sartori et al database

In the present study we discuss the Baetidae diversity of the differentbiogeographic realms with their rate of endemism and biogeographic affinities Wealso try to show the present state of knowledge of the different faunas and tohighlight those requiring the most attention

Palearctic realm

With 212 species the Palearctic has a high specific diversity (Table 1 Figure 1) Thisdiversity reflects the good knowledge of a great part of the realm rather than a higherdiversity than in other parts of the world The single genus Baetis itself representsabout half of the species part of them having a restricted distribution Most of the 17genera present a wide distribution including other realms The carnivorousRaptobaetopus and the rheophilous Acentrella are the only genera occurring in thePalearctic realm that show important adaptations all other genera are collector-gathers and have no or few adaptations to environmental factors

Four of the eight species of mayflies described by Linnaeus were from thePalearctic realm In the middle of the nineteenth century a few species weredescribed from Western Europe (Rambur 1842 Pictet 1843ndash1845) The contribution

Table 1 Generic and specific diversity by realm Endemicity of taxa endemic from thedifferent realm

No genera Endemicity No species Endemicity

Palearctic 17 12 212 98Afrotropics 40 82 190 100Oriental 23 38 124 99Neotropical 27 66 210 92Neartic 20 25 135 84Australasian 12 50 40 98

Aquatic Insects 43

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of Eaton to the knowledge of European Baetidae was essential in the description ofnew species and a new genus including the various monographs on mayflies hepublished and also with his comprehensive study of the mayflies of England (Eaton1870 Eaton 1871 Eaton 1883ndash1888) His work was outstanding due to the precisionof his drawings and the accuracy of the description He was also the first to have aconcept of the mayfly systematics which is still largely accurate

At the beginning of the twentieth century Bengtsson described several taxa basedon material from Scandinavia (Bengtsson 1912 1914 1917) Some of them werefound to have a much broader distribution in the West Palearctic Mainly based onsingle specimens pinned in different museums Navas described dozens of newspecies from Europe China and Japan (Navas 1911 1913 1917 1931b) Only a fewof them are still considered as valid most of them have never been mentioned sincethe original description Most species described after 1950 from Western Europerepresent alpine or insular vicariants (Thomas and Gazagnes 1984 Thomas 1986Thomas and Soldan 1987 Soldan and Godunko 2006) Several species weredescribed from East Europe (Bogoesco and Tabacaru 1957) and from East Palearctic(Kazlauskas 1963) The fauna of the Himalayas was subject to some attention butmore studies are required (Kapur and Kripalani 1963 Dubey 1971) The Japanesefauna encompasses 11 genera and 39 species most species are endemic to thearchipelago (Fujitani 2008) A great part of them were described before 1950 but themain contribution was made by Gose (1980) with the description of 14 speciesFujitani presented revisions of the Japanese Baetidae and described the larval stagefor several species (Fujitani et al 2003a 2003b) The Baetidae from Korea are alsoquite well-known but mainly included species with a wide distribution (Park et al1996 Bae 1997 Bae and Park 1998) In China several taxa in most of the families ofEphemeroptera were recently described or revised (Zhou et al 2000 Zhou andZheng 2001 Zhou and Braasch 2003 Zhou and Peters 2003 Zhou and Zheng 20032004) but nothing has been undertaken concerning the Baetidae

The study of North African fauna is relatively recent The Moroccan andAlgerian faunas possess several endemic species and show close relationships withWest Palearctic fauna mainly through the Iberian Peninsula (Soldan and Thomas1983a 1983b Thomas and Gagneur 1994 Thomas 1998) Afrotropical affinities arelimited to the single genus Cheleocloeon (Soldan and Thomas 1985) Levant and the

Figure 1 Specific and generic diversity in Palearctic realm Square total number of species(left scale) Triangle total number of genera (right scale)

44 J-L Gattolliat and C Nieto

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Arabian Peninsula have been also subject to attention recently as for North Africathis fauna possesses endemic species but a generic composition mainly similar to theEuropean one (Thomas and Dia 1984 1985 Thomas and Sartori 1989)

Afrotropical realm

The Afrotropical fauna of Baetidae presently encompasses 40 genera and 190 species(Figure 2) they represent about half of the generic and specific diversity of Africanmayflies All of the species and 82 of the genera of Baetidae are endemics from thisrealm the non-endemic genera possess a wide distribution including at least thePalearctic and Oriental realms (Baetis Cloeon and Nigrobaetis for example) TheAfrotropics clearly represent an important centre of diversity for the Baetidae

The distribution of some species covers a great part of the Afrotropics or even inthe case of Cloeon smaeleni the whole realm (Gattolliat and Rabeantoandro 2002)The specific distribution is sometimes greatly underestimated in part due to the lowdegree of knowledge of some faunas but also due to the description of the sametaxon under various names in different areas (Gattolliat unpublished data) Due topeculiar ecological conditions or geographic isolation some taxa have a restricteddistribution Madagascar the Drakensberg and Cape area in South Africa as well asthe volcanic mountain ranges in East and Central Africa are probably the main areasof microendemism

Ulmer (1909) described the first species of Afrotropical Baetidae fromMadagascar and the Comoros Subsequently a few other European researchersdescribed new taxa based on very restricted samples by generalist collectors whichended in European museums (Esben-Petersen 1913 Lestage 1918 Navas 19301931a) Barnard was the first true African ephemeropterist he published the firstmonograph on South African mayflies mainly from Western Cape (Barnard 1932)He was also one of the first who paid attention to the larval stage by rearing mayfliesto obtain larvandashadult association This pioneer study was followed by Crass (1947)and Agnew (1961 1963) contributing to the knowledge of the South African faunaAfter these three studies this fauna already comprised 26 species of Baetidae(compared to the 34 presently known) Most of them were described on a large seriesof specimens from different localities usually both at larval and imaginal stages

Figure 2 Specific and generic diversity in Afrotropical realm Square total number ofspecies (left scale) Triangle total number of genera (right scale)

Aquatic Insects 45

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Kimmins (1956) described a few species from Uganda Demoulin (1964a 1964b1965 1966 1967 1968) reported several species of Baetidae collected duringvarious expeditions In most cases he described them completely but refrained fromnaming them In 1970 he published the first catalogue of the African mayfly faunaincluding all the citations from the Afrotropical area (Demoulin 1970) Based onmore than 20000 imagines Kopelke (1980) described 12 new species from theKalengo mountains near Lake Kivu in Zaire This study was only based on theimaginal stage and the larva-imago association was subsequently possible only for afew species

Gillies contributed greatly to the knowledge of African Baetidae He first workedin East Africa mainly in Tanzania describing several species from this area (Gillies1985 1988 1990 1991a 1991b 1994 1998 Gillies and Wuillot 1997) In the 1970sand 1980s the French ORSTOM team collected aquatic macroinvertebratesrepeatedly in about 100 localities mainly in Guinea Ivory Coast Mali and SenegalThese collections constituted part of an important onchocerciasis controlprogramme (Leveque et al 2003) Elouard and Wuillot in close collaborationwith Gillies investigated this fauna and they described seven genera and about 20species from this area (Gillies 1990 Elouard and Hideux 1991 Wuillot and Gillies1993 1994)

The first mention of Baetidae in Madagascar was provided by Ulmer (1909)Until the middle of the 1990s little work on Malagasy Baetidae had been done atthat time eight species were described but only four of them are now considered asvalid (Gattolliat and Sartori 2003) In less than 10 years 19 genera and 51 specieswere described or reported from this island highlighting the great diversity of thisfauna this faunistic and systematic work was largely based on the material collectedby J-M Elouard and his Malagasy team (Elouard and Gibon 2001) To illustratethe originality and the highly adaptive nature of this fauna we can mention the threeendemic carnivorous genera (Gattolliat and Sartori 2001) the larva of Edmulmeatusgrandis feeding only on vascular plants (Gattolliat and Sartori 2003) and theadaptation for scraping algae from rocks occurring independently in several genera(Lugo-Ortiz and McCafferty 1997 Gattolliat and Sartori 2000 Gattolliat 2001a2001b 2002a 2002b 2004) Compared to other areas of the Afrotropics theMalagasy fauna can be considered as well known even if some systematics problemsneed to be solved and a few new species and genera still await description (Gattolliatand Sartori 2003 Gattolliat et al 2008)

Part of the Arabian Peninsula is also considered as belonging to the Afrotropicalrealm Despite a great part of this area being desert a limited but original faunaoccurs there (Gattolliat and Sartori 2008) This fauna shows direct influence fromboth the Palearctic and Afrotropical realms

In the 1980s and 1990s Waltz and McCafferty then Lugo-Ortiz and McCaffertyrevised the systematics of the Baetidae at the generic level They establishedrespectively three and 14 new genera (Lugo-Ortiz and McCafferty 1999b) It clearlyrepresented an important improvement of our knowledge of Afrotropical Baetidaebut still more systematic work is required before having a complete understanding ofthis fauna In the series lsquolsquoGuides to the Freshwater Invertebrates of SouthernAfricarsquorsquo Barber-James and Lugo-Ortiz (2003) offered a key to the generic level forlarvae of all the Afrotropical mayflies distribution diagnosis and ecology of eachgenus are discussed and an updated version of the checklist is available online(Barber-James 2007)

46 J-L Gattolliat and C Nieto

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We can consider that the faunas of South West and East Africa as well as ofMadagascar are relatively well known In all of these regions there are mostcertainly several species and genera that remain unknown because of restricteddistribution or unusual habitat The fauna of the whole Atlantic side of Africa (fromCameroon to Angola) remains almost completely unknown This fauna probablyencompasses some taxa with a broad distribution like Centroptiloides bifasciata orOphelmatostoma camerunse but it certainly contains several new taxa The area ofthe Ruwenzori mountain range is also poorly known and probably possesses a highdegree of endemism Almost no report of Baetidae is available from NortheasternAfrica north to Kenya

Oriental realm

The first species of Baetidae from the Oriental realm were described from SriLanka and India (Hagen 1858 Eaton 1883ndash1888) (Figure 3) In his work on themayflies of the Sunda Islands Ulmer (1939) provided the first comprehensivereview of an Oriental fauna with complete and accurate descriptions of new speciesat both larval and imaginal stages Kimmins (1947) and Gillies (1949) bothcontributed to the knowledge of the Baetidae from India by respectively describingthree and 11 species from the imaginal stage Apart from these works the Orientalfauna remained very poorly known until the 1980s Muller-Liebenau describedabout 60 new species from the larval stage (about 50 of the species presentlyknown from this area) and six new genera (Muller-Liebenau 1980a 1980b 19811982a 1982b 1982c 1983 1984a 1984b 1985 Muller-Liebenau and Heard 1979Muller-Liebenau and Morihara 1982 Muller-Liebenau and Hubbard 1985) Shemainly described species from Sri Lanka Malaysia and the Philippines Her workconstituted the major contribution to the knowledge of Oriental fauna Waltz andMcCafferty helped to clarify the supra-specific classification by establishing newgenera and complexes of genera (Waltz and McCafferty 1987a 1987c Waltz et al1994)

The Baetidae from Taiwan were the subject of an extensive study allowing thedescription of 14 species (Kang et al 1994 Kang and Yang 1996) The fauna fromrunning waters can be considered as well known but the fauna from still and

Figure 3 Specific and generic diversity in Oriental realm Square total number of species(left scale) Triangle total number of genera (right scale)

Aquatic Insects 47

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standing water remained unknown The fauna of Hong Kong has recently beensubject to close attention with the description of five new species (Tong and Dudgeon2000 2003)

Although some islands like Sri Lanka Taiwan or Hong Kong were the subject ofimportant studies the majority of the Oriental realm remains virtually unknownFor example only provisional checklists of Baetidae at generic level are available forVietnam and Thailand India especially the southern part of this subcontinentprobably possesses a much more diversified fauna than the scattered reports seem toindicate There are also almost no reports from countries with potentially highdiversity such as Cambodia or Myanmar If we consider that in a 85 km2 area inKalimantan (Borneo) intensive sampling allowed the collection of 12 genera ofBaetidae five of them probably being new to science (Sartori et al 2003) we caneasily imagine that only a tiny part of Oriental fauna is presently known Mostfaunas are too poorly known to correctly estimate the distribution of the differentspecimens but preliminary results based on our own collections seem to indicate thatat least some species show a wide distribution including a great part of South EastAsia It is extremely difficult to estimate the specific and generic diversity of theOriental realm even for restricted regions it is probably not overstated to considerthat the species presently known represent less than a half or even a third of the realdiversity

Neotropical realm

With a total surface of 19 million of km2 the Neotropical realm is the fourthbiogeographical region The Baetidae there now encompass 210 species and 27genera (Table 1) The first species of Baetidae from the Neotropics were describedby Pictet (1843ndash1845) Eaton (1871 1883ndash1888) and Weyenbergh (1883)(Figure 4) As for the Afrotropics the descriptions were based on restrictedcollections of imagines deposited in European museums Navas was the mostprolific author unfortunately he described the same species several times underdifferent names he described 26 species of Callibaetis but only eight are presentlyconsidering as valid (Domınguez et al 2006) Most of the species described bythese first authors have been transferred to new genera or synonymised

Figure 4 Specific and generic diversity in Neotropical realm Square total number of species(left scale) Triangle total number of genera (right scale)

48 J-L Gattolliat and C Nieto

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Traver and Edmunds (1968) greatly improved the knowledge of the highlydiversified genus Camelobaetidius Mol (1986) established the only carnivorousgenus for this region Harpagobaetis Lugo-Ortiz and McCafferty (1995 1996a1998c 1999e) described 12 new genera from the Neotropics other generahave been recently established by Nieto (2003b) and McCafferty and Baumgardner(2003) Finally Domınguez et al (2006) published the first comprehensivereview of all South America species of Ephemeroptera with a chapter on thefamily Baetidae including a key illustrations distribution and ecology for eachtaxon

The isolation of this continent for millions of years is one of the mostimportant reasons why 23 of the Neotropical genera and more than 90 of thespecies are endemics from this realm Only seven genera are shared with theNearctic region with a generally higher specific richness in the Neotropics(Apobaetis Baetodes Callibaetis Camelobaetidius and Paracloeodes) only Amer-icabaetis and Baetis are more diversified in the Nearctic (McCafferty 1998) SeveralSouth American species were originally described in the genus Baetis andPseudocloeon but they were all subsequently transferred to other genera such asCamelobaetidius Cloeodes Fallceon and Moribaetis (Lugo-Ortiz and McCafferty1999d McCafferty 2000) Baetis magnus is the only true Baetis which is present inthe Neotropical realm its distribution encompasses a great part of CentralAmerica and the South of USA Baetis is present therefore in all the realmsFallceon certainly also has a Panamerican distribution the generic attribution ofF candidus from Taiwan is clearly a mistake and consequently Fallceon is notpresent in the Oriental realm Cloeodes is in fact the only genus present in SouthAmerica with a wide distribution as species attributed to this genus have beenreported from the whole Pantropical area Most genera possessing an almostworldwide distribution such as Cloeon or LabiobaetisPseudocloeon are onlymissing in the Neotropics this absence also greatly contributes to the originality ofthe Neotropical fauna

Several Neotropical taxa present peculiar adaptations Aturbina is the only genusof the Baetidae without turbinate eyes Callibaetis is one of the few viviparous generain the Ephemeroptera Chane is a spectacular genus with labium and maxillaeextremely well developed for collecting and filtering (Nieto 2003b) presentingsimilarities with the Afroptropical genus Ophelmatostoma (Gillies et al 1990) Mayo(1973) described the first species collected in a waterfall Baetis ellenae latertransferred to Mayobaetis (Waltz and McCafferty 1985) Cloeodes hydation istolerant to short periods of habitat desiccation this species occurs in smalltemporary rock pools (Nolte et al 1996)

Knowledge of the Baetidae from this region has been greatly improved in the lastfew decades new species and genera were described (Dominique et al 2002 Nieto2003a 2003b 2004a 2004b Thomas and Peru 2003) and other genera were reviewed(Nieto 2004c Nieto and Salles 2006 Nieto and Richard 2008) However there arestill regions or even countries with poorly known faunas such as Bolivia Chile PeruEcuador Uruguay and Venezuela these areas almost certainly possess someendemic taxa which remain to be found Recent samplings in Brazilian Amazoniaallowed the discovery of several new species (Salles and Batista 2004 Sallesand Francischetti 2004 Salles et al 2004 2005 Salles 2007 Salles and Polegatto2008) but a great part of this fauna remains unknown (Salles personalcommunication)

Aquatic Insects 49

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Nearctic realm

Being the second largest biogeographic region (229 million km2) the Nearctic regionencompasses only 20 described genera (Table 1) The rate of endemism is also lowOnly five genera have a distribution restricted to this area (25) Seven genera areshared with the Neotropical region other genera present a distributionincluding the Palearctic andor Oriental areas Most genera are collector-gatherersor collector-scrapers no carnivorous Baetidae have been recorded from this regionFrom the 135 Nearctic species 114 are endemic (84) four Holarctic and 17Panamerican

The first ephemeropterologists working in this region were Hagen (1861) Walsh(1863a 1863b) and Eaton (1869 1870 1871 1881 1883ndash1888) Banks (1900 19141918) described eight species of Callibaetis five of which are still valid With 42species McDunnough was the main knowledgeable contributor to Nearctic Baetidae(McDunnough 1922 1923 1924 1925 1926 1929 1931 1932 1936 1939)(Figure 5)

The first accurate revision of all the species from North America was made byNeedham et al (1935) At that time the concept of Baetidae encompassed severaltaxonomic groups that are presently considered as separate families Edmundsrsquo workon mayflies was significant also in this area He proposed one of the first checklists ofmayflies from North America and Mexico (Edmunds and Allen 1957a) Hiscontribution to the biogeography and evolution of Ephemeroptera is still largelyaccurate (Edmunds 1972 1975) In 1976 he published in collaboration with Jensenand Berner a book with all the species known at that time from North and CentralAmerica (Edmunds et al 1976) In the Baetidae he described and reassigned severalspecies (Edmunds 1954 Edmunds and Allen 1957b)

In the Nearctic region contemporary authors such as Jacobus (Jacobus andMcCafferty 2005a 2005b 2006) Lugo-Ortiz and McCafferty (Lugo-Ortiz andMcCafferty 1998d) Waltz (Waltz and McCafferty 1986 1987b 1989) and Wiersema(Wiersema 2000 Wiersema and McCafferty 2004 Wiersema et al 2004) alsocontributed to the advanced knowledge of this fauna by the description of generaand species as well as several revisions with generic reassignments They sustain anactualised web page with a checklist and general information of the mayflies ofNorth and Central America (McCafferty 2009)

Figure 5 Specific and generic diversity in Nearctic realm Square total number of species(left scale) Triangle total number of genera (right scale)

50 J-L Gattolliat and C Nieto

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After 150 years of intensive works the systematics of Baetidae can be consideredas relatively well known even if some taxomomic problems are still not completelysolved The validity of some species needs to be reviewed a few of them have neverbeen mentioned since their original description

Australasian realm

The Australasian realm presently encompasses 40 known species all of themendemic except for Cloeon virens which is also distributed in the Oriental realmTwelve genera occur in this area only six of them being endemic (50) (Table 1)The non-endemic genera are also distributed in the Oriental region The presence ofsome typical Palearctic genera such as Centroptilum in Australasia is doubtful andshould be clarified in the near future

Offadens soror was the first species described from this region by Ulmer in 1908Following Ulmer Tillyard and Harker provided the only significant works onAustralian Baetidae (Tillyard 1936 Harker 1950 1954 1957) and until recently(Figure 6) Suter greatly improved the knowledge of the Ephemeroptera fauna in theAustralian region providing the first key for the larval stage of mayflies (Suter 1979)At that time the Baetidae encompassed only 12 species and five genera andinterestingly Bungona was the only genus which was endemic from this area all theother Australian species being attributed to Palearctic or Holarctic genera Later onSuter described new species and redescribed some genera (Suter 1986 2000 2001Suter and Pearson 2001) He also provided a key specifically for the larvae ofBaetidae which included several undescribed species and genera (Suter 1997)

As in other biogeographic regions Lugo-Ortiz and McCafferty played animportant role by describing several species and four endemic genera from thisregion (Lugo-Ortiz and McCafferty 1998e 1999a 1999c) and by reporting generasuch as Cloeodes or LabiobaetisPseudocloeon (Lugo-Ortiz and McCafferty 1998bLugo-Ortiz et al 1999) They also described several new taxa from New Guinea(Lugo-Ortiz and McCafferty 1999c Lugo-Ortiz et al 1999) Baetidae were one of thetwo families of mayflies reported from the Fiji Islands although it was only apreliminary study the diversity was relatively high with 12 species and three genera(Flowers 1990)

Figure 6 Specific and generic diversity in Australasian realm Square total number ofspecies (left scale) Triangle total number of genera (right scale)

Aquatic Insects 51

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Although the predaceous habit is very unusual in Baetidae two of the 12 generaof this region are carnivorous Echinobaetis from Sulawesi (Mol 1989) andMystaxyops from New Guinea (McCafferty and Sun 2005)

New Zealand is the only important landmass where mayflies are present andBaetidae have never been found This gap remains an enigma These islands are quiteisolated and possess endemic families such as Siphlaenigmatidae and Rallidentidaebut also the widespread family Leptophlebiidae The Baetidae were also consideredto be absent from New Caledonia (Peters 2001) However in material recentlycollected in standing waters half a dozen larvae of Cloeon were included(unpublished data) Because of the absence of Baetidae in previous intensivesamplings and the high power of dispersion of Cloeon it seems likely that a recentcolonisation of the island may be related to human activity If the Leptophlebiidaewere able to remain established on these islands after the break-off of the Gondwanaor to colonise the islands afterwards why were the Baetidae not able to do the sameThe question remains open

Despite recent improvements the fauna of this region remains insufficientlyknown in Australia and even more so on other islands and archipelagos ForAustralia a recent estimation established that only about half of the Australianspecies of Baetidae are described and probably some new genera still need to beestablished (Jeff Webb personal communication)

The biogeographic affinities of this region need to be analysed in detail All thenon-endemic genera are also distributed in the Oriental region they are mainlydiversified in the North East of Australasia and had probably dispersed step by stepthrough South East Asia into Australia where their distribution is limited to theNorth (Lugo-Ortiz and McCafferty 1999b) The genus Cloeodes clearly connects thisarea with the history of Gondwana and the continents placed in the SouthHemisphere Endemic genera such Edmundsiops Offadens and Bungona may berelated to South American taxa (Jeff Webb personal communication)

Conclusion

Since the establishment of the Baetidae in 1815 their knowledge has been hugelyincreased due to the efforts of many authors around the world In 1999 Lugo-Ortizand McCafferty provided the first worldwide synthesis of this family They estimatedthe diversity of this family at 650 species and 87 genera (Lugo-Ortiz and McCafferty1999b) Ten years later the number of species has increased by almost one third toreach 894 species During these 10 years 142 new species were described mainlyfrom South America Madagascar and West Africa It means that the increase of thetotal number of species is mainly due to the description of new species but Lugo-Ortiz and McCafferty had also underestimated the number of species of Baetidae byabout 100 species During these 10 years 17 genera were described and four weresynonymised

Baetidae are mostly diversified and possess a much higher rate of genericendemism in the Tropics and Southern Hemisphere than in the NorthernHemisphere (Barber-James et al 2008) The Afrotropics and Neotropics possessby far the most diversified faunas Despite common Gondwanian origins thesefaunas have almost no affinities The Palearctic fauna shares most genera with theOriental andor Afrotropical faunas as well as the Nearctic fauna Stepwisecolonisation could have occurred through the Palearctic region Central America

52 J-L Gattolliat and C Nieto

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constitutes a bridge between North and South Americas and is under the influence ofboth areas (McCafferty 1998) The affinities of the Nearctic and Neotropical faunasare mainly due to the presence of North American taxa in Central America TheOriental fauna is still only partially known regions of biogeographic importance andwith a high potential diversity such as the Indian subcontinent and South East Asiaremain almost completely unknown Local studies clearly indicate that the diversityis quite high but the distribution of the taxa remains unknown The Oriental realm istherefore probably the most promising and challenging area Even if the globaldiversity is probably lower in Australasia the lack of knowledge of the fauna isobvious especially in the tropical part of this area such as New Guinea

The knowledge on systematics and faunistic composition are the first inevitablesteps to understand and preserve a natural system Most of the remaining unknownor poorly known faunas are under a direct human threat This underlines theurgency of this work before these faunas are definitively lost this is especially true inmost of the tropical rain forest areas where a primordial aquatic fauna is still presentFor areas with sufficient knowledge it is also the duty of the systematists to offer thenecessary tools for a correct identification of the taxa We can only encourageeveryone to make our knowledge accessible to others scientists especially by creatingkeys and checklists at different systematic levels

A comprehensive reconstruction of the phylogeny of the Baetidae is also highlynecessary Such a reconstruction should be based on modern tools and include bothmolecular and morphological characters It is the only way to solve contradictorysuprageneric relationships Such studies would allow a more accurate understandingof the relationships between the different areas in a global biogeographic approachThey would help to understand the historical influence of Gondwana and Laurasiaon the mayfly fauna

Acknowledgements

We want to express our appreciation to Michel Sartori (Museum of Zoology LausanneSwitzerland) and Helen Barber-James (Albany Museum Grahamstown South Africa) foruseful comments on this paper and great help in the conception of the Baetidae databaseAline Pasche (Museum of Zoology Lausanne Switzerland) offered technical supportFinancial support from the Argentine National Council of Scientific and TechnologicalResearch (CONICET) and ANPCYT (Pict 524-Pict 528) to which the junior author belongsare acknowledged The electronic database lsquolsquoEphemeroptera of the Worldrsquorsquo of Nikita Kluge(httpwwwinsectabiopuruzEph-sppContentshtm) was of great help for cross-checkingof our own database

References

Agnew JD (1961) lsquoNew Baetidae (Ephem) from South Africarsquo Novos Taxa Entomologicos25 3ndash18

Agnew JD (1963) lsquoNew South African records of imperfectly known Baetidae (Ephem)rsquoHydrobiologia 22 41ndash46

Bae YJ (1997) lsquoTaxonomy of Cloeon and Procloeon (Ephemeroptera Baetidae) in KorearsquoThe Korean Journal of Systematics Biology 13 303ndash314

Bae YJ and Park SY (1998) lsquoAlainites Baetis Labiobaetis and Nigrobaetis(Ephemeroptera Baetidae) in Korearsquo The Korean Journal of Systematics Biology 141ndash12

Banks N (1900) lsquoNew genera and species of Nearctic Neuropteroid Insectsrsquo Transactions ofthe American Entomological Society 26 239ndash259

Banks N (1914) lsquoNew neuropteroid insects native and exoticrsquo Proceedings of the Academy ofNatural Sciences of Philadelphia 66 608ndash632

Aquatic Insects 53

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of Eaton to the knowledge of European Baetidae was essential in the description ofnew species and a new genus including the various monographs on mayflies hepublished and also with his comprehensive study of the mayflies of England (Eaton1870 Eaton 1871 Eaton 1883ndash1888) His work was outstanding due to the precisionof his drawings and the accuracy of the description He was also the first to have aconcept of the mayfly systematics which is still largely accurate

At the beginning of the twentieth century Bengtsson described several taxa basedon material from Scandinavia (Bengtsson 1912 1914 1917) Some of them werefound to have a much broader distribution in the West Palearctic Mainly based onsingle specimens pinned in different museums Navas described dozens of newspecies from Europe China and Japan (Navas 1911 1913 1917 1931b) Only a fewof them are still considered as valid most of them have never been mentioned sincethe original description Most species described after 1950 from Western Europerepresent alpine or insular vicariants (Thomas and Gazagnes 1984 Thomas 1986Thomas and Soldan 1987 Soldan and Godunko 2006) Several species weredescribed from East Europe (Bogoesco and Tabacaru 1957) and from East Palearctic(Kazlauskas 1963) The fauna of the Himalayas was subject to some attention butmore studies are required (Kapur and Kripalani 1963 Dubey 1971) The Japanesefauna encompasses 11 genera and 39 species most species are endemic to thearchipelago (Fujitani 2008) A great part of them were described before 1950 but themain contribution was made by Gose (1980) with the description of 14 speciesFujitani presented revisions of the Japanese Baetidae and described the larval stagefor several species (Fujitani et al 2003a 2003b) The Baetidae from Korea are alsoquite well-known but mainly included species with a wide distribution (Park et al1996 Bae 1997 Bae and Park 1998) In China several taxa in most of the families ofEphemeroptera were recently described or revised (Zhou et al 2000 Zhou andZheng 2001 Zhou and Braasch 2003 Zhou and Peters 2003 Zhou and Zheng 20032004) but nothing has been undertaken concerning the Baetidae

The study of North African fauna is relatively recent The Moroccan andAlgerian faunas possess several endemic species and show close relationships withWest Palearctic fauna mainly through the Iberian Peninsula (Soldan and Thomas1983a 1983b Thomas and Gagneur 1994 Thomas 1998) Afrotropical affinities arelimited to the single genus Cheleocloeon (Soldan and Thomas 1985) Levant and the

Figure 1 Specific and generic diversity in Palearctic realm Square total number of species(left scale) Triangle total number of genera (right scale)

44 J-L Gattolliat and C Nieto

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Arabian Peninsula have been also subject to attention recently as for North Africathis fauna possesses endemic species but a generic composition mainly similar to theEuropean one (Thomas and Dia 1984 1985 Thomas and Sartori 1989)

Afrotropical realm

The Afrotropical fauna of Baetidae presently encompasses 40 genera and 190 species(Figure 2) they represent about half of the generic and specific diversity of Africanmayflies All of the species and 82 of the genera of Baetidae are endemics from thisrealm the non-endemic genera possess a wide distribution including at least thePalearctic and Oriental realms (Baetis Cloeon and Nigrobaetis for example) TheAfrotropics clearly represent an important centre of diversity for the Baetidae

The distribution of some species covers a great part of the Afrotropics or even inthe case of Cloeon smaeleni the whole realm (Gattolliat and Rabeantoandro 2002)The specific distribution is sometimes greatly underestimated in part due to the lowdegree of knowledge of some faunas but also due to the description of the sametaxon under various names in different areas (Gattolliat unpublished data) Due topeculiar ecological conditions or geographic isolation some taxa have a restricteddistribution Madagascar the Drakensberg and Cape area in South Africa as well asthe volcanic mountain ranges in East and Central Africa are probably the main areasof microendemism

Ulmer (1909) described the first species of Afrotropical Baetidae fromMadagascar and the Comoros Subsequently a few other European researchersdescribed new taxa based on very restricted samples by generalist collectors whichended in European museums (Esben-Petersen 1913 Lestage 1918 Navas 19301931a) Barnard was the first true African ephemeropterist he published the firstmonograph on South African mayflies mainly from Western Cape (Barnard 1932)He was also one of the first who paid attention to the larval stage by rearing mayfliesto obtain larvandashadult association This pioneer study was followed by Crass (1947)and Agnew (1961 1963) contributing to the knowledge of the South African faunaAfter these three studies this fauna already comprised 26 species of Baetidae(compared to the 34 presently known) Most of them were described on a large seriesof specimens from different localities usually both at larval and imaginal stages

Figure 2 Specific and generic diversity in Afrotropical realm Square total number ofspecies (left scale) Triangle total number of genera (right scale)

Aquatic Insects 45

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Kimmins (1956) described a few species from Uganda Demoulin (1964a 1964b1965 1966 1967 1968) reported several species of Baetidae collected duringvarious expeditions In most cases he described them completely but refrained fromnaming them In 1970 he published the first catalogue of the African mayfly faunaincluding all the citations from the Afrotropical area (Demoulin 1970) Based onmore than 20000 imagines Kopelke (1980) described 12 new species from theKalengo mountains near Lake Kivu in Zaire This study was only based on theimaginal stage and the larva-imago association was subsequently possible only for afew species

Gillies contributed greatly to the knowledge of African Baetidae He first workedin East Africa mainly in Tanzania describing several species from this area (Gillies1985 1988 1990 1991a 1991b 1994 1998 Gillies and Wuillot 1997) In the 1970sand 1980s the French ORSTOM team collected aquatic macroinvertebratesrepeatedly in about 100 localities mainly in Guinea Ivory Coast Mali and SenegalThese collections constituted part of an important onchocerciasis controlprogramme (Leveque et al 2003) Elouard and Wuillot in close collaborationwith Gillies investigated this fauna and they described seven genera and about 20species from this area (Gillies 1990 Elouard and Hideux 1991 Wuillot and Gillies1993 1994)

The first mention of Baetidae in Madagascar was provided by Ulmer (1909)Until the middle of the 1990s little work on Malagasy Baetidae had been done atthat time eight species were described but only four of them are now considered asvalid (Gattolliat and Sartori 2003) In less than 10 years 19 genera and 51 specieswere described or reported from this island highlighting the great diversity of thisfauna this faunistic and systematic work was largely based on the material collectedby J-M Elouard and his Malagasy team (Elouard and Gibon 2001) To illustratethe originality and the highly adaptive nature of this fauna we can mention the threeendemic carnivorous genera (Gattolliat and Sartori 2001) the larva of Edmulmeatusgrandis feeding only on vascular plants (Gattolliat and Sartori 2003) and theadaptation for scraping algae from rocks occurring independently in several genera(Lugo-Ortiz and McCafferty 1997 Gattolliat and Sartori 2000 Gattolliat 2001a2001b 2002a 2002b 2004) Compared to other areas of the Afrotropics theMalagasy fauna can be considered as well known even if some systematics problemsneed to be solved and a few new species and genera still await description (Gattolliatand Sartori 2003 Gattolliat et al 2008)

Part of the Arabian Peninsula is also considered as belonging to the Afrotropicalrealm Despite a great part of this area being desert a limited but original faunaoccurs there (Gattolliat and Sartori 2008) This fauna shows direct influence fromboth the Palearctic and Afrotropical realms

In the 1980s and 1990s Waltz and McCafferty then Lugo-Ortiz and McCaffertyrevised the systematics of the Baetidae at the generic level They establishedrespectively three and 14 new genera (Lugo-Ortiz and McCafferty 1999b) It clearlyrepresented an important improvement of our knowledge of Afrotropical Baetidaebut still more systematic work is required before having a complete understanding ofthis fauna In the series lsquolsquoGuides to the Freshwater Invertebrates of SouthernAfricarsquorsquo Barber-James and Lugo-Ortiz (2003) offered a key to the generic level forlarvae of all the Afrotropical mayflies distribution diagnosis and ecology of eachgenus are discussed and an updated version of the checklist is available online(Barber-James 2007)

46 J-L Gattolliat and C Nieto

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We can consider that the faunas of South West and East Africa as well as ofMadagascar are relatively well known In all of these regions there are mostcertainly several species and genera that remain unknown because of restricteddistribution or unusual habitat The fauna of the whole Atlantic side of Africa (fromCameroon to Angola) remains almost completely unknown This fauna probablyencompasses some taxa with a broad distribution like Centroptiloides bifasciata orOphelmatostoma camerunse but it certainly contains several new taxa The area ofthe Ruwenzori mountain range is also poorly known and probably possesses a highdegree of endemism Almost no report of Baetidae is available from NortheasternAfrica north to Kenya

Oriental realm

The first species of Baetidae from the Oriental realm were described from SriLanka and India (Hagen 1858 Eaton 1883ndash1888) (Figure 3) In his work on themayflies of the Sunda Islands Ulmer (1939) provided the first comprehensivereview of an Oriental fauna with complete and accurate descriptions of new speciesat both larval and imaginal stages Kimmins (1947) and Gillies (1949) bothcontributed to the knowledge of the Baetidae from India by respectively describingthree and 11 species from the imaginal stage Apart from these works the Orientalfauna remained very poorly known until the 1980s Muller-Liebenau describedabout 60 new species from the larval stage (about 50 of the species presentlyknown from this area) and six new genera (Muller-Liebenau 1980a 1980b 19811982a 1982b 1982c 1983 1984a 1984b 1985 Muller-Liebenau and Heard 1979Muller-Liebenau and Morihara 1982 Muller-Liebenau and Hubbard 1985) Shemainly described species from Sri Lanka Malaysia and the Philippines Her workconstituted the major contribution to the knowledge of Oriental fauna Waltz andMcCafferty helped to clarify the supra-specific classification by establishing newgenera and complexes of genera (Waltz and McCafferty 1987a 1987c Waltz et al1994)

The Baetidae from Taiwan were the subject of an extensive study allowing thedescription of 14 species (Kang et al 1994 Kang and Yang 1996) The fauna fromrunning waters can be considered as well known but the fauna from still and

Figure 3 Specific and generic diversity in Oriental realm Square total number of species(left scale) Triangle total number of genera (right scale)

Aquatic Insects 47

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standing water remained unknown The fauna of Hong Kong has recently beensubject to close attention with the description of five new species (Tong and Dudgeon2000 2003)

Although some islands like Sri Lanka Taiwan or Hong Kong were the subject ofimportant studies the majority of the Oriental realm remains virtually unknownFor example only provisional checklists of Baetidae at generic level are available forVietnam and Thailand India especially the southern part of this subcontinentprobably possesses a much more diversified fauna than the scattered reports seem toindicate There are also almost no reports from countries with potentially highdiversity such as Cambodia or Myanmar If we consider that in a 85 km2 area inKalimantan (Borneo) intensive sampling allowed the collection of 12 genera ofBaetidae five of them probably being new to science (Sartori et al 2003) we caneasily imagine that only a tiny part of Oriental fauna is presently known Mostfaunas are too poorly known to correctly estimate the distribution of the differentspecimens but preliminary results based on our own collections seem to indicate thatat least some species show a wide distribution including a great part of South EastAsia It is extremely difficult to estimate the specific and generic diversity of theOriental realm even for restricted regions it is probably not overstated to considerthat the species presently known represent less than a half or even a third of the realdiversity

Neotropical realm

With a total surface of 19 million of km2 the Neotropical realm is the fourthbiogeographical region The Baetidae there now encompass 210 species and 27genera (Table 1) The first species of Baetidae from the Neotropics were describedby Pictet (1843ndash1845) Eaton (1871 1883ndash1888) and Weyenbergh (1883)(Figure 4) As for the Afrotropics the descriptions were based on restrictedcollections of imagines deposited in European museums Navas was the mostprolific author unfortunately he described the same species several times underdifferent names he described 26 species of Callibaetis but only eight are presentlyconsidering as valid (Domınguez et al 2006) Most of the species described bythese first authors have been transferred to new genera or synonymised

Figure 4 Specific and generic diversity in Neotropical realm Square total number of species(left scale) Triangle total number of genera (right scale)

48 J-L Gattolliat and C Nieto

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Traver and Edmunds (1968) greatly improved the knowledge of the highlydiversified genus Camelobaetidius Mol (1986) established the only carnivorousgenus for this region Harpagobaetis Lugo-Ortiz and McCafferty (1995 1996a1998c 1999e) described 12 new genera from the Neotropics other generahave been recently established by Nieto (2003b) and McCafferty and Baumgardner(2003) Finally Domınguez et al (2006) published the first comprehensivereview of all South America species of Ephemeroptera with a chapter on thefamily Baetidae including a key illustrations distribution and ecology for eachtaxon

The isolation of this continent for millions of years is one of the mostimportant reasons why 23 of the Neotropical genera and more than 90 of thespecies are endemics from this realm Only seven genera are shared with theNearctic region with a generally higher specific richness in the Neotropics(Apobaetis Baetodes Callibaetis Camelobaetidius and Paracloeodes) only Amer-icabaetis and Baetis are more diversified in the Nearctic (McCafferty 1998) SeveralSouth American species were originally described in the genus Baetis andPseudocloeon but they were all subsequently transferred to other genera such asCamelobaetidius Cloeodes Fallceon and Moribaetis (Lugo-Ortiz and McCafferty1999d McCafferty 2000) Baetis magnus is the only true Baetis which is present inthe Neotropical realm its distribution encompasses a great part of CentralAmerica and the South of USA Baetis is present therefore in all the realmsFallceon certainly also has a Panamerican distribution the generic attribution ofF candidus from Taiwan is clearly a mistake and consequently Fallceon is notpresent in the Oriental realm Cloeodes is in fact the only genus present in SouthAmerica with a wide distribution as species attributed to this genus have beenreported from the whole Pantropical area Most genera possessing an almostworldwide distribution such as Cloeon or LabiobaetisPseudocloeon are onlymissing in the Neotropics this absence also greatly contributes to the originality ofthe Neotropical fauna

Several Neotropical taxa present peculiar adaptations Aturbina is the only genusof the Baetidae without turbinate eyes Callibaetis is one of the few viviparous generain the Ephemeroptera Chane is a spectacular genus with labium and maxillaeextremely well developed for collecting and filtering (Nieto 2003b) presentingsimilarities with the Afroptropical genus Ophelmatostoma (Gillies et al 1990) Mayo(1973) described the first species collected in a waterfall Baetis ellenae latertransferred to Mayobaetis (Waltz and McCafferty 1985) Cloeodes hydation istolerant to short periods of habitat desiccation this species occurs in smalltemporary rock pools (Nolte et al 1996)

Knowledge of the Baetidae from this region has been greatly improved in the lastfew decades new species and genera were described (Dominique et al 2002 Nieto2003a 2003b 2004a 2004b Thomas and Peru 2003) and other genera were reviewed(Nieto 2004c Nieto and Salles 2006 Nieto and Richard 2008) However there arestill regions or even countries with poorly known faunas such as Bolivia Chile PeruEcuador Uruguay and Venezuela these areas almost certainly possess someendemic taxa which remain to be found Recent samplings in Brazilian Amazoniaallowed the discovery of several new species (Salles and Batista 2004 Sallesand Francischetti 2004 Salles et al 2004 2005 Salles 2007 Salles and Polegatto2008) but a great part of this fauna remains unknown (Salles personalcommunication)

Aquatic Insects 49

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Nearctic realm

Being the second largest biogeographic region (229 million km2) the Nearctic regionencompasses only 20 described genera (Table 1) The rate of endemism is also lowOnly five genera have a distribution restricted to this area (25) Seven genera areshared with the Neotropical region other genera present a distributionincluding the Palearctic andor Oriental areas Most genera are collector-gatherersor collector-scrapers no carnivorous Baetidae have been recorded from this regionFrom the 135 Nearctic species 114 are endemic (84) four Holarctic and 17Panamerican

The first ephemeropterologists working in this region were Hagen (1861) Walsh(1863a 1863b) and Eaton (1869 1870 1871 1881 1883ndash1888) Banks (1900 19141918) described eight species of Callibaetis five of which are still valid With 42species McDunnough was the main knowledgeable contributor to Nearctic Baetidae(McDunnough 1922 1923 1924 1925 1926 1929 1931 1932 1936 1939)(Figure 5)

The first accurate revision of all the species from North America was made byNeedham et al (1935) At that time the concept of Baetidae encompassed severaltaxonomic groups that are presently considered as separate families Edmundsrsquo workon mayflies was significant also in this area He proposed one of the first checklists ofmayflies from North America and Mexico (Edmunds and Allen 1957a) Hiscontribution to the biogeography and evolution of Ephemeroptera is still largelyaccurate (Edmunds 1972 1975) In 1976 he published in collaboration with Jensenand Berner a book with all the species known at that time from North and CentralAmerica (Edmunds et al 1976) In the Baetidae he described and reassigned severalspecies (Edmunds 1954 Edmunds and Allen 1957b)

In the Nearctic region contemporary authors such as Jacobus (Jacobus andMcCafferty 2005a 2005b 2006) Lugo-Ortiz and McCafferty (Lugo-Ortiz andMcCafferty 1998d) Waltz (Waltz and McCafferty 1986 1987b 1989) and Wiersema(Wiersema 2000 Wiersema and McCafferty 2004 Wiersema et al 2004) alsocontributed to the advanced knowledge of this fauna by the description of generaand species as well as several revisions with generic reassignments They sustain anactualised web page with a checklist and general information of the mayflies ofNorth and Central America (McCafferty 2009)

Figure 5 Specific and generic diversity in Nearctic realm Square total number of species(left scale) Triangle total number of genera (right scale)

50 J-L Gattolliat and C Nieto

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After 150 years of intensive works the systematics of Baetidae can be consideredas relatively well known even if some taxomomic problems are still not completelysolved The validity of some species needs to be reviewed a few of them have neverbeen mentioned since their original description

Australasian realm

The Australasian realm presently encompasses 40 known species all of themendemic except for Cloeon virens which is also distributed in the Oriental realmTwelve genera occur in this area only six of them being endemic (50) (Table 1)The non-endemic genera are also distributed in the Oriental region The presence ofsome typical Palearctic genera such as Centroptilum in Australasia is doubtful andshould be clarified in the near future

Offadens soror was the first species described from this region by Ulmer in 1908Following Ulmer Tillyard and Harker provided the only significant works onAustralian Baetidae (Tillyard 1936 Harker 1950 1954 1957) and until recently(Figure 6) Suter greatly improved the knowledge of the Ephemeroptera fauna in theAustralian region providing the first key for the larval stage of mayflies (Suter 1979)At that time the Baetidae encompassed only 12 species and five genera andinterestingly Bungona was the only genus which was endemic from this area all theother Australian species being attributed to Palearctic or Holarctic genera Later onSuter described new species and redescribed some genera (Suter 1986 2000 2001Suter and Pearson 2001) He also provided a key specifically for the larvae ofBaetidae which included several undescribed species and genera (Suter 1997)

As in other biogeographic regions Lugo-Ortiz and McCafferty played animportant role by describing several species and four endemic genera from thisregion (Lugo-Ortiz and McCafferty 1998e 1999a 1999c) and by reporting generasuch as Cloeodes or LabiobaetisPseudocloeon (Lugo-Ortiz and McCafferty 1998bLugo-Ortiz et al 1999) They also described several new taxa from New Guinea(Lugo-Ortiz and McCafferty 1999c Lugo-Ortiz et al 1999) Baetidae were one of thetwo families of mayflies reported from the Fiji Islands although it was only apreliminary study the diversity was relatively high with 12 species and three genera(Flowers 1990)

Figure 6 Specific and generic diversity in Australasian realm Square total number ofspecies (left scale) Triangle total number of genera (right scale)

Aquatic Insects 51

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Although the predaceous habit is very unusual in Baetidae two of the 12 generaof this region are carnivorous Echinobaetis from Sulawesi (Mol 1989) andMystaxyops from New Guinea (McCafferty and Sun 2005)

New Zealand is the only important landmass where mayflies are present andBaetidae have never been found This gap remains an enigma These islands are quiteisolated and possess endemic families such as Siphlaenigmatidae and Rallidentidaebut also the widespread family Leptophlebiidae The Baetidae were also consideredto be absent from New Caledonia (Peters 2001) However in material recentlycollected in standing waters half a dozen larvae of Cloeon were included(unpublished data) Because of the absence of Baetidae in previous intensivesamplings and the high power of dispersion of Cloeon it seems likely that a recentcolonisation of the island may be related to human activity If the Leptophlebiidaewere able to remain established on these islands after the break-off of the Gondwanaor to colonise the islands afterwards why were the Baetidae not able to do the sameThe question remains open

Despite recent improvements the fauna of this region remains insufficientlyknown in Australia and even more so on other islands and archipelagos ForAustralia a recent estimation established that only about half of the Australianspecies of Baetidae are described and probably some new genera still need to beestablished (Jeff Webb personal communication)

The biogeographic affinities of this region need to be analysed in detail All thenon-endemic genera are also distributed in the Oriental region they are mainlydiversified in the North East of Australasia and had probably dispersed step by stepthrough South East Asia into Australia where their distribution is limited to theNorth (Lugo-Ortiz and McCafferty 1999b) The genus Cloeodes clearly connects thisarea with the history of Gondwana and the continents placed in the SouthHemisphere Endemic genera such Edmundsiops Offadens and Bungona may berelated to South American taxa (Jeff Webb personal communication)

Conclusion

Since the establishment of the Baetidae in 1815 their knowledge has been hugelyincreased due to the efforts of many authors around the world In 1999 Lugo-Ortizand McCafferty provided the first worldwide synthesis of this family They estimatedthe diversity of this family at 650 species and 87 genera (Lugo-Ortiz and McCafferty1999b) Ten years later the number of species has increased by almost one third toreach 894 species During these 10 years 142 new species were described mainlyfrom South America Madagascar and West Africa It means that the increase of thetotal number of species is mainly due to the description of new species but Lugo-Ortiz and McCafferty had also underestimated the number of species of Baetidae byabout 100 species During these 10 years 17 genera were described and four weresynonymised

Baetidae are mostly diversified and possess a much higher rate of genericendemism in the Tropics and Southern Hemisphere than in the NorthernHemisphere (Barber-James et al 2008) The Afrotropics and Neotropics possessby far the most diversified faunas Despite common Gondwanian origins thesefaunas have almost no affinities The Palearctic fauna shares most genera with theOriental andor Afrotropical faunas as well as the Nearctic fauna Stepwisecolonisation could have occurred through the Palearctic region Central America

52 J-L Gattolliat and C Nieto

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constitutes a bridge between North and South Americas and is under the influence ofboth areas (McCafferty 1998) The affinities of the Nearctic and Neotropical faunasare mainly due to the presence of North American taxa in Central America TheOriental fauna is still only partially known regions of biogeographic importance andwith a high potential diversity such as the Indian subcontinent and South East Asiaremain almost completely unknown Local studies clearly indicate that the diversityis quite high but the distribution of the taxa remains unknown The Oriental realm istherefore probably the most promising and challenging area Even if the globaldiversity is probably lower in Australasia the lack of knowledge of the fauna isobvious especially in the tropical part of this area such as New Guinea

The knowledge on systematics and faunistic composition are the first inevitablesteps to understand and preserve a natural system Most of the remaining unknownor poorly known faunas are under a direct human threat This underlines theurgency of this work before these faunas are definitively lost this is especially true inmost of the tropical rain forest areas where a primordial aquatic fauna is still presentFor areas with sufficient knowledge it is also the duty of the systematists to offer thenecessary tools for a correct identification of the taxa We can only encourageeveryone to make our knowledge accessible to others scientists especially by creatingkeys and checklists at different systematic levels

A comprehensive reconstruction of the phylogeny of the Baetidae is also highlynecessary Such a reconstruction should be based on modern tools and include bothmolecular and morphological characters It is the only way to solve contradictorysuprageneric relationships Such studies would allow a more accurate understandingof the relationships between the different areas in a global biogeographic approachThey would help to understand the historical influence of Gondwana and Laurasiaon the mayfly fauna

Acknowledgements

We want to express our appreciation to Michel Sartori (Museum of Zoology LausanneSwitzerland) and Helen Barber-James (Albany Museum Grahamstown South Africa) foruseful comments on this paper and great help in the conception of the Baetidae databaseAline Pasche (Museum of Zoology Lausanne Switzerland) offered technical supportFinancial support from the Argentine National Council of Scientific and TechnologicalResearch (CONICET) and ANPCYT (Pict 524-Pict 528) to which the junior author belongsare acknowledged The electronic database lsquolsquoEphemeroptera of the Worldrsquorsquo of Nikita Kluge(httpwwwinsectabiopuruzEph-sppContentshtm) was of great help for cross-checkingof our own database

References

Agnew JD (1961) lsquoNew Baetidae (Ephem) from South Africarsquo Novos Taxa Entomologicos25 3ndash18

Agnew JD (1963) lsquoNew South African records of imperfectly known Baetidae (Ephem)rsquoHydrobiologia 22 41ndash46

Bae YJ (1997) lsquoTaxonomy of Cloeon and Procloeon (Ephemeroptera Baetidae) in KorearsquoThe Korean Journal of Systematics Biology 13 303ndash314

Bae YJ and Park SY (1998) lsquoAlainites Baetis Labiobaetis and Nigrobaetis(Ephemeroptera Baetidae) in Korearsquo The Korean Journal of Systematics Biology 141ndash12

Banks N (1900) lsquoNew genera and species of Nearctic Neuropteroid Insectsrsquo Transactions ofthe American Entomological Society 26 239ndash259

Banks N (1914) lsquoNew neuropteroid insects native and exoticrsquo Proceedings of the Academy ofNatural Sciences of Philadelphia 66 608ndash632

Aquatic Insects 53

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Arabian Peninsula have been also subject to attention recently as for North Africathis fauna possesses endemic species but a generic composition mainly similar to theEuropean one (Thomas and Dia 1984 1985 Thomas and Sartori 1989)

Afrotropical realm

The Afrotropical fauna of Baetidae presently encompasses 40 genera and 190 species(Figure 2) they represent about half of the generic and specific diversity of Africanmayflies All of the species and 82 of the genera of Baetidae are endemics from thisrealm the non-endemic genera possess a wide distribution including at least thePalearctic and Oriental realms (Baetis Cloeon and Nigrobaetis for example) TheAfrotropics clearly represent an important centre of diversity for the Baetidae

The distribution of some species covers a great part of the Afrotropics or even inthe case of Cloeon smaeleni the whole realm (Gattolliat and Rabeantoandro 2002)The specific distribution is sometimes greatly underestimated in part due to the lowdegree of knowledge of some faunas but also due to the description of the sametaxon under various names in different areas (Gattolliat unpublished data) Due topeculiar ecological conditions or geographic isolation some taxa have a restricteddistribution Madagascar the Drakensberg and Cape area in South Africa as well asthe volcanic mountain ranges in East and Central Africa are probably the main areasof microendemism

Ulmer (1909) described the first species of Afrotropical Baetidae fromMadagascar and the Comoros Subsequently a few other European researchersdescribed new taxa based on very restricted samples by generalist collectors whichended in European museums (Esben-Petersen 1913 Lestage 1918 Navas 19301931a) Barnard was the first true African ephemeropterist he published the firstmonograph on South African mayflies mainly from Western Cape (Barnard 1932)He was also one of the first who paid attention to the larval stage by rearing mayfliesto obtain larvandashadult association This pioneer study was followed by Crass (1947)and Agnew (1961 1963) contributing to the knowledge of the South African faunaAfter these three studies this fauna already comprised 26 species of Baetidae(compared to the 34 presently known) Most of them were described on a large seriesof specimens from different localities usually both at larval and imaginal stages

Figure 2 Specific and generic diversity in Afrotropical realm Square total number ofspecies (left scale) Triangle total number of genera (right scale)

Aquatic Insects 45

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Kimmins (1956) described a few species from Uganda Demoulin (1964a 1964b1965 1966 1967 1968) reported several species of Baetidae collected duringvarious expeditions In most cases he described them completely but refrained fromnaming them In 1970 he published the first catalogue of the African mayfly faunaincluding all the citations from the Afrotropical area (Demoulin 1970) Based onmore than 20000 imagines Kopelke (1980) described 12 new species from theKalengo mountains near Lake Kivu in Zaire This study was only based on theimaginal stage and the larva-imago association was subsequently possible only for afew species

Gillies contributed greatly to the knowledge of African Baetidae He first workedin East Africa mainly in Tanzania describing several species from this area (Gillies1985 1988 1990 1991a 1991b 1994 1998 Gillies and Wuillot 1997) In the 1970sand 1980s the French ORSTOM team collected aquatic macroinvertebratesrepeatedly in about 100 localities mainly in Guinea Ivory Coast Mali and SenegalThese collections constituted part of an important onchocerciasis controlprogramme (Leveque et al 2003) Elouard and Wuillot in close collaborationwith Gillies investigated this fauna and they described seven genera and about 20species from this area (Gillies 1990 Elouard and Hideux 1991 Wuillot and Gillies1993 1994)

The first mention of Baetidae in Madagascar was provided by Ulmer (1909)Until the middle of the 1990s little work on Malagasy Baetidae had been done atthat time eight species were described but only four of them are now considered asvalid (Gattolliat and Sartori 2003) In less than 10 years 19 genera and 51 specieswere described or reported from this island highlighting the great diversity of thisfauna this faunistic and systematic work was largely based on the material collectedby J-M Elouard and his Malagasy team (Elouard and Gibon 2001) To illustratethe originality and the highly adaptive nature of this fauna we can mention the threeendemic carnivorous genera (Gattolliat and Sartori 2001) the larva of Edmulmeatusgrandis feeding only on vascular plants (Gattolliat and Sartori 2003) and theadaptation for scraping algae from rocks occurring independently in several genera(Lugo-Ortiz and McCafferty 1997 Gattolliat and Sartori 2000 Gattolliat 2001a2001b 2002a 2002b 2004) Compared to other areas of the Afrotropics theMalagasy fauna can be considered as well known even if some systematics problemsneed to be solved and a few new species and genera still await description (Gattolliatand Sartori 2003 Gattolliat et al 2008)

Part of the Arabian Peninsula is also considered as belonging to the Afrotropicalrealm Despite a great part of this area being desert a limited but original faunaoccurs there (Gattolliat and Sartori 2008) This fauna shows direct influence fromboth the Palearctic and Afrotropical realms

In the 1980s and 1990s Waltz and McCafferty then Lugo-Ortiz and McCaffertyrevised the systematics of the Baetidae at the generic level They establishedrespectively three and 14 new genera (Lugo-Ortiz and McCafferty 1999b) It clearlyrepresented an important improvement of our knowledge of Afrotropical Baetidaebut still more systematic work is required before having a complete understanding ofthis fauna In the series lsquolsquoGuides to the Freshwater Invertebrates of SouthernAfricarsquorsquo Barber-James and Lugo-Ortiz (2003) offered a key to the generic level forlarvae of all the Afrotropical mayflies distribution diagnosis and ecology of eachgenus are discussed and an updated version of the checklist is available online(Barber-James 2007)

46 J-L Gattolliat and C Nieto

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We can consider that the faunas of South West and East Africa as well as ofMadagascar are relatively well known In all of these regions there are mostcertainly several species and genera that remain unknown because of restricteddistribution or unusual habitat The fauna of the whole Atlantic side of Africa (fromCameroon to Angola) remains almost completely unknown This fauna probablyencompasses some taxa with a broad distribution like Centroptiloides bifasciata orOphelmatostoma camerunse but it certainly contains several new taxa The area ofthe Ruwenzori mountain range is also poorly known and probably possesses a highdegree of endemism Almost no report of Baetidae is available from NortheasternAfrica north to Kenya

Oriental realm

The first species of Baetidae from the Oriental realm were described from SriLanka and India (Hagen 1858 Eaton 1883ndash1888) (Figure 3) In his work on themayflies of the Sunda Islands Ulmer (1939) provided the first comprehensivereview of an Oriental fauna with complete and accurate descriptions of new speciesat both larval and imaginal stages Kimmins (1947) and Gillies (1949) bothcontributed to the knowledge of the Baetidae from India by respectively describingthree and 11 species from the imaginal stage Apart from these works the Orientalfauna remained very poorly known until the 1980s Muller-Liebenau describedabout 60 new species from the larval stage (about 50 of the species presentlyknown from this area) and six new genera (Muller-Liebenau 1980a 1980b 19811982a 1982b 1982c 1983 1984a 1984b 1985 Muller-Liebenau and Heard 1979Muller-Liebenau and Morihara 1982 Muller-Liebenau and Hubbard 1985) Shemainly described species from Sri Lanka Malaysia and the Philippines Her workconstituted the major contribution to the knowledge of Oriental fauna Waltz andMcCafferty helped to clarify the supra-specific classification by establishing newgenera and complexes of genera (Waltz and McCafferty 1987a 1987c Waltz et al1994)

The Baetidae from Taiwan were the subject of an extensive study allowing thedescription of 14 species (Kang et al 1994 Kang and Yang 1996) The fauna fromrunning waters can be considered as well known but the fauna from still and

Figure 3 Specific and generic diversity in Oriental realm Square total number of species(left scale) Triangle total number of genera (right scale)

Aquatic Insects 47

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standing water remained unknown The fauna of Hong Kong has recently beensubject to close attention with the description of five new species (Tong and Dudgeon2000 2003)

Although some islands like Sri Lanka Taiwan or Hong Kong were the subject ofimportant studies the majority of the Oriental realm remains virtually unknownFor example only provisional checklists of Baetidae at generic level are available forVietnam and Thailand India especially the southern part of this subcontinentprobably possesses a much more diversified fauna than the scattered reports seem toindicate There are also almost no reports from countries with potentially highdiversity such as Cambodia or Myanmar If we consider that in a 85 km2 area inKalimantan (Borneo) intensive sampling allowed the collection of 12 genera ofBaetidae five of them probably being new to science (Sartori et al 2003) we caneasily imagine that only a tiny part of Oriental fauna is presently known Mostfaunas are too poorly known to correctly estimate the distribution of the differentspecimens but preliminary results based on our own collections seem to indicate thatat least some species show a wide distribution including a great part of South EastAsia It is extremely difficult to estimate the specific and generic diversity of theOriental realm even for restricted regions it is probably not overstated to considerthat the species presently known represent less than a half or even a third of the realdiversity

Neotropical realm

With a total surface of 19 million of km2 the Neotropical realm is the fourthbiogeographical region The Baetidae there now encompass 210 species and 27genera (Table 1) The first species of Baetidae from the Neotropics were describedby Pictet (1843ndash1845) Eaton (1871 1883ndash1888) and Weyenbergh (1883)(Figure 4) As for the Afrotropics the descriptions were based on restrictedcollections of imagines deposited in European museums Navas was the mostprolific author unfortunately he described the same species several times underdifferent names he described 26 species of Callibaetis but only eight are presentlyconsidering as valid (Domınguez et al 2006) Most of the species described bythese first authors have been transferred to new genera or synonymised

Figure 4 Specific and generic diversity in Neotropical realm Square total number of species(left scale) Triangle total number of genera (right scale)

48 J-L Gattolliat and C Nieto

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Traver and Edmunds (1968) greatly improved the knowledge of the highlydiversified genus Camelobaetidius Mol (1986) established the only carnivorousgenus for this region Harpagobaetis Lugo-Ortiz and McCafferty (1995 1996a1998c 1999e) described 12 new genera from the Neotropics other generahave been recently established by Nieto (2003b) and McCafferty and Baumgardner(2003) Finally Domınguez et al (2006) published the first comprehensivereview of all South America species of Ephemeroptera with a chapter on thefamily Baetidae including a key illustrations distribution and ecology for eachtaxon

The isolation of this continent for millions of years is one of the mostimportant reasons why 23 of the Neotropical genera and more than 90 of thespecies are endemics from this realm Only seven genera are shared with theNearctic region with a generally higher specific richness in the Neotropics(Apobaetis Baetodes Callibaetis Camelobaetidius and Paracloeodes) only Amer-icabaetis and Baetis are more diversified in the Nearctic (McCafferty 1998) SeveralSouth American species were originally described in the genus Baetis andPseudocloeon but they were all subsequently transferred to other genera such asCamelobaetidius Cloeodes Fallceon and Moribaetis (Lugo-Ortiz and McCafferty1999d McCafferty 2000) Baetis magnus is the only true Baetis which is present inthe Neotropical realm its distribution encompasses a great part of CentralAmerica and the South of USA Baetis is present therefore in all the realmsFallceon certainly also has a Panamerican distribution the generic attribution ofF candidus from Taiwan is clearly a mistake and consequently Fallceon is notpresent in the Oriental realm Cloeodes is in fact the only genus present in SouthAmerica with a wide distribution as species attributed to this genus have beenreported from the whole Pantropical area Most genera possessing an almostworldwide distribution such as Cloeon or LabiobaetisPseudocloeon are onlymissing in the Neotropics this absence also greatly contributes to the originality ofthe Neotropical fauna

Several Neotropical taxa present peculiar adaptations Aturbina is the only genusof the Baetidae without turbinate eyes Callibaetis is one of the few viviparous generain the Ephemeroptera Chane is a spectacular genus with labium and maxillaeextremely well developed for collecting and filtering (Nieto 2003b) presentingsimilarities with the Afroptropical genus Ophelmatostoma (Gillies et al 1990) Mayo(1973) described the first species collected in a waterfall Baetis ellenae latertransferred to Mayobaetis (Waltz and McCafferty 1985) Cloeodes hydation istolerant to short periods of habitat desiccation this species occurs in smalltemporary rock pools (Nolte et al 1996)

Knowledge of the Baetidae from this region has been greatly improved in the lastfew decades new species and genera were described (Dominique et al 2002 Nieto2003a 2003b 2004a 2004b Thomas and Peru 2003) and other genera were reviewed(Nieto 2004c Nieto and Salles 2006 Nieto and Richard 2008) However there arestill regions or even countries with poorly known faunas such as Bolivia Chile PeruEcuador Uruguay and Venezuela these areas almost certainly possess someendemic taxa which remain to be found Recent samplings in Brazilian Amazoniaallowed the discovery of several new species (Salles and Batista 2004 Sallesand Francischetti 2004 Salles et al 2004 2005 Salles 2007 Salles and Polegatto2008) but a great part of this fauna remains unknown (Salles personalcommunication)

Aquatic Insects 49

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Nearctic realm

Being the second largest biogeographic region (229 million km2) the Nearctic regionencompasses only 20 described genera (Table 1) The rate of endemism is also lowOnly five genera have a distribution restricted to this area (25) Seven genera areshared with the Neotropical region other genera present a distributionincluding the Palearctic andor Oriental areas Most genera are collector-gatherersor collector-scrapers no carnivorous Baetidae have been recorded from this regionFrom the 135 Nearctic species 114 are endemic (84) four Holarctic and 17Panamerican

The first ephemeropterologists working in this region were Hagen (1861) Walsh(1863a 1863b) and Eaton (1869 1870 1871 1881 1883ndash1888) Banks (1900 19141918) described eight species of Callibaetis five of which are still valid With 42species McDunnough was the main knowledgeable contributor to Nearctic Baetidae(McDunnough 1922 1923 1924 1925 1926 1929 1931 1932 1936 1939)(Figure 5)

The first accurate revision of all the species from North America was made byNeedham et al (1935) At that time the concept of Baetidae encompassed severaltaxonomic groups that are presently considered as separate families Edmundsrsquo workon mayflies was significant also in this area He proposed one of the first checklists ofmayflies from North America and Mexico (Edmunds and Allen 1957a) Hiscontribution to the biogeography and evolution of Ephemeroptera is still largelyaccurate (Edmunds 1972 1975) In 1976 he published in collaboration with Jensenand Berner a book with all the species known at that time from North and CentralAmerica (Edmunds et al 1976) In the Baetidae he described and reassigned severalspecies (Edmunds 1954 Edmunds and Allen 1957b)

In the Nearctic region contemporary authors such as Jacobus (Jacobus andMcCafferty 2005a 2005b 2006) Lugo-Ortiz and McCafferty (Lugo-Ortiz andMcCafferty 1998d) Waltz (Waltz and McCafferty 1986 1987b 1989) and Wiersema(Wiersema 2000 Wiersema and McCafferty 2004 Wiersema et al 2004) alsocontributed to the advanced knowledge of this fauna by the description of generaand species as well as several revisions with generic reassignments They sustain anactualised web page with a checklist and general information of the mayflies ofNorth and Central America (McCafferty 2009)

Figure 5 Specific and generic diversity in Nearctic realm Square total number of species(left scale) Triangle total number of genera (right scale)

50 J-L Gattolliat and C Nieto

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After 150 years of intensive works the systematics of Baetidae can be consideredas relatively well known even if some taxomomic problems are still not completelysolved The validity of some species needs to be reviewed a few of them have neverbeen mentioned since their original description

Australasian realm

The Australasian realm presently encompasses 40 known species all of themendemic except for Cloeon virens which is also distributed in the Oriental realmTwelve genera occur in this area only six of them being endemic (50) (Table 1)The non-endemic genera are also distributed in the Oriental region The presence ofsome typical Palearctic genera such as Centroptilum in Australasia is doubtful andshould be clarified in the near future

Offadens soror was the first species described from this region by Ulmer in 1908Following Ulmer Tillyard and Harker provided the only significant works onAustralian Baetidae (Tillyard 1936 Harker 1950 1954 1957) and until recently(Figure 6) Suter greatly improved the knowledge of the Ephemeroptera fauna in theAustralian region providing the first key for the larval stage of mayflies (Suter 1979)At that time the Baetidae encompassed only 12 species and five genera andinterestingly Bungona was the only genus which was endemic from this area all theother Australian species being attributed to Palearctic or Holarctic genera Later onSuter described new species and redescribed some genera (Suter 1986 2000 2001Suter and Pearson 2001) He also provided a key specifically for the larvae ofBaetidae which included several undescribed species and genera (Suter 1997)

As in other biogeographic regions Lugo-Ortiz and McCafferty played animportant role by describing several species and four endemic genera from thisregion (Lugo-Ortiz and McCafferty 1998e 1999a 1999c) and by reporting generasuch as Cloeodes or LabiobaetisPseudocloeon (Lugo-Ortiz and McCafferty 1998bLugo-Ortiz et al 1999) They also described several new taxa from New Guinea(Lugo-Ortiz and McCafferty 1999c Lugo-Ortiz et al 1999) Baetidae were one of thetwo families of mayflies reported from the Fiji Islands although it was only apreliminary study the diversity was relatively high with 12 species and three genera(Flowers 1990)

Figure 6 Specific and generic diversity in Australasian realm Square total number ofspecies (left scale) Triangle total number of genera (right scale)

Aquatic Insects 51

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Although the predaceous habit is very unusual in Baetidae two of the 12 generaof this region are carnivorous Echinobaetis from Sulawesi (Mol 1989) andMystaxyops from New Guinea (McCafferty and Sun 2005)

New Zealand is the only important landmass where mayflies are present andBaetidae have never been found This gap remains an enigma These islands are quiteisolated and possess endemic families such as Siphlaenigmatidae and Rallidentidaebut also the widespread family Leptophlebiidae The Baetidae were also consideredto be absent from New Caledonia (Peters 2001) However in material recentlycollected in standing waters half a dozen larvae of Cloeon were included(unpublished data) Because of the absence of Baetidae in previous intensivesamplings and the high power of dispersion of Cloeon it seems likely that a recentcolonisation of the island may be related to human activity If the Leptophlebiidaewere able to remain established on these islands after the break-off of the Gondwanaor to colonise the islands afterwards why were the Baetidae not able to do the sameThe question remains open

Despite recent improvements the fauna of this region remains insufficientlyknown in Australia and even more so on other islands and archipelagos ForAustralia a recent estimation established that only about half of the Australianspecies of Baetidae are described and probably some new genera still need to beestablished (Jeff Webb personal communication)

The biogeographic affinities of this region need to be analysed in detail All thenon-endemic genera are also distributed in the Oriental region they are mainlydiversified in the North East of Australasia and had probably dispersed step by stepthrough South East Asia into Australia where their distribution is limited to theNorth (Lugo-Ortiz and McCafferty 1999b) The genus Cloeodes clearly connects thisarea with the history of Gondwana and the continents placed in the SouthHemisphere Endemic genera such Edmundsiops Offadens and Bungona may berelated to South American taxa (Jeff Webb personal communication)

Conclusion

Since the establishment of the Baetidae in 1815 their knowledge has been hugelyincreased due to the efforts of many authors around the world In 1999 Lugo-Ortizand McCafferty provided the first worldwide synthesis of this family They estimatedthe diversity of this family at 650 species and 87 genera (Lugo-Ortiz and McCafferty1999b) Ten years later the number of species has increased by almost one third toreach 894 species During these 10 years 142 new species were described mainlyfrom South America Madagascar and West Africa It means that the increase of thetotal number of species is mainly due to the description of new species but Lugo-Ortiz and McCafferty had also underestimated the number of species of Baetidae byabout 100 species During these 10 years 17 genera were described and four weresynonymised

Baetidae are mostly diversified and possess a much higher rate of genericendemism in the Tropics and Southern Hemisphere than in the NorthernHemisphere (Barber-James et al 2008) The Afrotropics and Neotropics possessby far the most diversified faunas Despite common Gondwanian origins thesefaunas have almost no affinities The Palearctic fauna shares most genera with theOriental andor Afrotropical faunas as well as the Nearctic fauna Stepwisecolonisation could have occurred through the Palearctic region Central America

52 J-L Gattolliat and C Nieto

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constitutes a bridge between North and South Americas and is under the influence ofboth areas (McCafferty 1998) The affinities of the Nearctic and Neotropical faunasare mainly due to the presence of North American taxa in Central America TheOriental fauna is still only partially known regions of biogeographic importance andwith a high potential diversity such as the Indian subcontinent and South East Asiaremain almost completely unknown Local studies clearly indicate that the diversityis quite high but the distribution of the taxa remains unknown The Oriental realm istherefore probably the most promising and challenging area Even if the globaldiversity is probably lower in Australasia the lack of knowledge of the fauna isobvious especially in the tropical part of this area such as New Guinea

The knowledge on systematics and faunistic composition are the first inevitablesteps to understand and preserve a natural system Most of the remaining unknownor poorly known faunas are under a direct human threat This underlines theurgency of this work before these faunas are definitively lost this is especially true inmost of the tropical rain forest areas where a primordial aquatic fauna is still presentFor areas with sufficient knowledge it is also the duty of the systematists to offer thenecessary tools for a correct identification of the taxa We can only encourageeveryone to make our knowledge accessible to others scientists especially by creatingkeys and checklists at different systematic levels

A comprehensive reconstruction of the phylogeny of the Baetidae is also highlynecessary Such a reconstruction should be based on modern tools and include bothmolecular and morphological characters It is the only way to solve contradictorysuprageneric relationships Such studies would allow a more accurate understandingof the relationships between the different areas in a global biogeographic approachThey would help to understand the historical influence of Gondwana and Laurasiaon the mayfly fauna

Acknowledgements

We want to express our appreciation to Michel Sartori (Museum of Zoology LausanneSwitzerland) and Helen Barber-James (Albany Museum Grahamstown South Africa) foruseful comments on this paper and great help in the conception of the Baetidae databaseAline Pasche (Museum of Zoology Lausanne Switzerland) offered technical supportFinancial support from the Argentine National Council of Scientific and TechnologicalResearch (CONICET) and ANPCYT (Pict 524-Pict 528) to which the junior author belongsare acknowledged The electronic database lsquolsquoEphemeroptera of the Worldrsquorsquo of Nikita Kluge(httpwwwinsectabiopuruzEph-sppContentshtm) was of great help for cross-checkingof our own database

References

Agnew JD (1961) lsquoNew Baetidae (Ephem) from South Africarsquo Novos Taxa Entomologicos25 3ndash18

Agnew JD (1963) lsquoNew South African records of imperfectly known Baetidae (Ephem)rsquoHydrobiologia 22 41ndash46

Bae YJ (1997) lsquoTaxonomy of Cloeon and Procloeon (Ephemeroptera Baetidae) in KorearsquoThe Korean Journal of Systematics Biology 13 303ndash314

Bae YJ and Park SY (1998) lsquoAlainites Baetis Labiobaetis and Nigrobaetis(Ephemeroptera Baetidae) in Korearsquo The Korean Journal of Systematics Biology 141ndash12

Banks N (1900) lsquoNew genera and species of Nearctic Neuropteroid Insectsrsquo Transactions ofthe American Entomological Society 26 239ndash259

Banks N (1914) lsquoNew neuropteroid insects native and exoticrsquo Proceedings of the Academy ofNatural Sciences of Philadelphia 66 608ndash632

Aquatic Insects 53

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Kimmins (1956) described a few species from Uganda Demoulin (1964a 1964b1965 1966 1967 1968) reported several species of Baetidae collected duringvarious expeditions In most cases he described them completely but refrained fromnaming them In 1970 he published the first catalogue of the African mayfly faunaincluding all the citations from the Afrotropical area (Demoulin 1970) Based onmore than 20000 imagines Kopelke (1980) described 12 new species from theKalengo mountains near Lake Kivu in Zaire This study was only based on theimaginal stage and the larva-imago association was subsequently possible only for afew species

Gillies contributed greatly to the knowledge of African Baetidae He first workedin East Africa mainly in Tanzania describing several species from this area (Gillies1985 1988 1990 1991a 1991b 1994 1998 Gillies and Wuillot 1997) In the 1970sand 1980s the French ORSTOM team collected aquatic macroinvertebratesrepeatedly in about 100 localities mainly in Guinea Ivory Coast Mali and SenegalThese collections constituted part of an important onchocerciasis controlprogramme (Leveque et al 2003) Elouard and Wuillot in close collaborationwith Gillies investigated this fauna and they described seven genera and about 20species from this area (Gillies 1990 Elouard and Hideux 1991 Wuillot and Gillies1993 1994)

The first mention of Baetidae in Madagascar was provided by Ulmer (1909)Until the middle of the 1990s little work on Malagasy Baetidae had been done atthat time eight species were described but only four of them are now considered asvalid (Gattolliat and Sartori 2003) In less than 10 years 19 genera and 51 specieswere described or reported from this island highlighting the great diversity of thisfauna this faunistic and systematic work was largely based on the material collectedby J-M Elouard and his Malagasy team (Elouard and Gibon 2001) To illustratethe originality and the highly adaptive nature of this fauna we can mention the threeendemic carnivorous genera (Gattolliat and Sartori 2001) the larva of Edmulmeatusgrandis feeding only on vascular plants (Gattolliat and Sartori 2003) and theadaptation for scraping algae from rocks occurring independently in several genera(Lugo-Ortiz and McCafferty 1997 Gattolliat and Sartori 2000 Gattolliat 2001a2001b 2002a 2002b 2004) Compared to other areas of the Afrotropics theMalagasy fauna can be considered as well known even if some systematics problemsneed to be solved and a few new species and genera still await description (Gattolliatand Sartori 2003 Gattolliat et al 2008)

Part of the Arabian Peninsula is also considered as belonging to the Afrotropicalrealm Despite a great part of this area being desert a limited but original faunaoccurs there (Gattolliat and Sartori 2008) This fauna shows direct influence fromboth the Palearctic and Afrotropical realms

In the 1980s and 1990s Waltz and McCafferty then Lugo-Ortiz and McCaffertyrevised the systematics of the Baetidae at the generic level They establishedrespectively three and 14 new genera (Lugo-Ortiz and McCafferty 1999b) It clearlyrepresented an important improvement of our knowledge of Afrotropical Baetidaebut still more systematic work is required before having a complete understanding ofthis fauna In the series lsquolsquoGuides to the Freshwater Invertebrates of SouthernAfricarsquorsquo Barber-James and Lugo-Ortiz (2003) offered a key to the generic level forlarvae of all the Afrotropical mayflies distribution diagnosis and ecology of eachgenus are discussed and an updated version of the checklist is available online(Barber-James 2007)

46 J-L Gattolliat and C Nieto

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13

We can consider that the faunas of South West and East Africa as well as ofMadagascar are relatively well known In all of these regions there are mostcertainly several species and genera that remain unknown because of restricteddistribution or unusual habitat The fauna of the whole Atlantic side of Africa (fromCameroon to Angola) remains almost completely unknown This fauna probablyencompasses some taxa with a broad distribution like Centroptiloides bifasciata orOphelmatostoma camerunse but it certainly contains several new taxa The area ofthe Ruwenzori mountain range is also poorly known and probably possesses a highdegree of endemism Almost no report of Baetidae is available from NortheasternAfrica north to Kenya

Oriental realm

The first species of Baetidae from the Oriental realm were described from SriLanka and India (Hagen 1858 Eaton 1883ndash1888) (Figure 3) In his work on themayflies of the Sunda Islands Ulmer (1939) provided the first comprehensivereview of an Oriental fauna with complete and accurate descriptions of new speciesat both larval and imaginal stages Kimmins (1947) and Gillies (1949) bothcontributed to the knowledge of the Baetidae from India by respectively describingthree and 11 species from the imaginal stage Apart from these works the Orientalfauna remained very poorly known until the 1980s Muller-Liebenau describedabout 60 new species from the larval stage (about 50 of the species presentlyknown from this area) and six new genera (Muller-Liebenau 1980a 1980b 19811982a 1982b 1982c 1983 1984a 1984b 1985 Muller-Liebenau and Heard 1979Muller-Liebenau and Morihara 1982 Muller-Liebenau and Hubbard 1985) Shemainly described species from Sri Lanka Malaysia and the Philippines Her workconstituted the major contribution to the knowledge of Oriental fauna Waltz andMcCafferty helped to clarify the supra-specific classification by establishing newgenera and complexes of genera (Waltz and McCafferty 1987a 1987c Waltz et al1994)

The Baetidae from Taiwan were the subject of an extensive study allowing thedescription of 14 species (Kang et al 1994 Kang and Yang 1996) The fauna fromrunning waters can be considered as well known but the fauna from still and

Figure 3 Specific and generic diversity in Oriental realm Square total number of species(left scale) Triangle total number of genera (right scale)

Aquatic Insects 47

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standing water remained unknown The fauna of Hong Kong has recently beensubject to close attention with the description of five new species (Tong and Dudgeon2000 2003)

Although some islands like Sri Lanka Taiwan or Hong Kong were the subject ofimportant studies the majority of the Oriental realm remains virtually unknownFor example only provisional checklists of Baetidae at generic level are available forVietnam and Thailand India especially the southern part of this subcontinentprobably possesses a much more diversified fauna than the scattered reports seem toindicate There are also almost no reports from countries with potentially highdiversity such as Cambodia or Myanmar If we consider that in a 85 km2 area inKalimantan (Borneo) intensive sampling allowed the collection of 12 genera ofBaetidae five of them probably being new to science (Sartori et al 2003) we caneasily imagine that only a tiny part of Oriental fauna is presently known Mostfaunas are too poorly known to correctly estimate the distribution of the differentspecimens but preliminary results based on our own collections seem to indicate thatat least some species show a wide distribution including a great part of South EastAsia It is extremely difficult to estimate the specific and generic diversity of theOriental realm even for restricted regions it is probably not overstated to considerthat the species presently known represent less than a half or even a third of the realdiversity

Neotropical realm

With a total surface of 19 million of km2 the Neotropical realm is the fourthbiogeographical region The Baetidae there now encompass 210 species and 27genera (Table 1) The first species of Baetidae from the Neotropics were describedby Pictet (1843ndash1845) Eaton (1871 1883ndash1888) and Weyenbergh (1883)(Figure 4) As for the Afrotropics the descriptions were based on restrictedcollections of imagines deposited in European museums Navas was the mostprolific author unfortunately he described the same species several times underdifferent names he described 26 species of Callibaetis but only eight are presentlyconsidering as valid (Domınguez et al 2006) Most of the species described bythese first authors have been transferred to new genera or synonymised

Figure 4 Specific and generic diversity in Neotropical realm Square total number of species(left scale) Triangle total number of genera (right scale)

48 J-L Gattolliat and C Nieto

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Sept

embe

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13

Traver and Edmunds (1968) greatly improved the knowledge of the highlydiversified genus Camelobaetidius Mol (1986) established the only carnivorousgenus for this region Harpagobaetis Lugo-Ortiz and McCafferty (1995 1996a1998c 1999e) described 12 new genera from the Neotropics other generahave been recently established by Nieto (2003b) and McCafferty and Baumgardner(2003) Finally Domınguez et al (2006) published the first comprehensivereview of all South America species of Ephemeroptera with a chapter on thefamily Baetidae including a key illustrations distribution and ecology for eachtaxon

The isolation of this continent for millions of years is one of the mostimportant reasons why 23 of the Neotropical genera and more than 90 of thespecies are endemics from this realm Only seven genera are shared with theNearctic region with a generally higher specific richness in the Neotropics(Apobaetis Baetodes Callibaetis Camelobaetidius and Paracloeodes) only Amer-icabaetis and Baetis are more diversified in the Nearctic (McCafferty 1998) SeveralSouth American species were originally described in the genus Baetis andPseudocloeon but they were all subsequently transferred to other genera such asCamelobaetidius Cloeodes Fallceon and Moribaetis (Lugo-Ortiz and McCafferty1999d McCafferty 2000) Baetis magnus is the only true Baetis which is present inthe Neotropical realm its distribution encompasses a great part of CentralAmerica and the South of USA Baetis is present therefore in all the realmsFallceon certainly also has a Panamerican distribution the generic attribution ofF candidus from Taiwan is clearly a mistake and consequently Fallceon is notpresent in the Oriental realm Cloeodes is in fact the only genus present in SouthAmerica with a wide distribution as species attributed to this genus have beenreported from the whole Pantropical area Most genera possessing an almostworldwide distribution such as Cloeon or LabiobaetisPseudocloeon are onlymissing in the Neotropics this absence also greatly contributes to the originality ofthe Neotropical fauna

Several Neotropical taxa present peculiar adaptations Aturbina is the only genusof the Baetidae without turbinate eyes Callibaetis is one of the few viviparous generain the Ephemeroptera Chane is a spectacular genus with labium and maxillaeextremely well developed for collecting and filtering (Nieto 2003b) presentingsimilarities with the Afroptropical genus Ophelmatostoma (Gillies et al 1990) Mayo(1973) described the first species collected in a waterfall Baetis ellenae latertransferred to Mayobaetis (Waltz and McCafferty 1985) Cloeodes hydation istolerant to short periods of habitat desiccation this species occurs in smalltemporary rock pools (Nolte et al 1996)

Knowledge of the Baetidae from this region has been greatly improved in the lastfew decades new species and genera were described (Dominique et al 2002 Nieto2003a 2003b 2004a 2004b Thomas and Peru 2003) and other genera were reviewed(Nieto 2004c Nieto and Salles 2006 Nieto and Richard 2008) However there arestill regions or even countries with poorly known faunas such as Bolivia Chile PeruEcuador Uruguay and Venezuela these areas almost certainly possess someendemic taxa which remain to be found Recent samplings in Brazilian Amazoniaallowed the discovery of several new species (Salles and Batista 2004 Sallesand Francischetti 2004 Salles et al 2004 2005 Salles 2007 Salles and Polegatto2008) but a great part of this fauna remains unknown (Salles personalcommunication)

Aquatic Insects 49

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Nearctic realm

Being the second largest biogeographic region (229 million km2) the Nearctic regionencompasses only 20 described genera (Table 1) The rate of endemism is also lowOnly five genera have a distribution restricted to this area (25) Seven genera areshared with the Neotropical region other genera present a distributionincluding the Palearctic andor Oriental areas Most genera are collector-gatherersor collector-scrapers no carnivorous Baetidae have been recorded from this regionFrom the 135 Nearctic species 114 are endemic (84) four Holarctic and 17Panamerican

The first ephemeropterologists working in this region were Hagen (1861) Walsh(1863a 1863b) and Eaton (1869 1870 1871 1881 1883ndash1888) Banks (1900 19141918) described eight species of Callibaetis five of which are still valid With 42species McDunnough was the main knowledgeable contributor to Nearctic Baetidae(McDunnough 1922 1923 1924 1925 1926 1929 1931 1932 1936 1939)(Figure 5)

The first accurate revision of all the species from North America was made byNeedham et al (1935) At that time the concept of Baetidae encompassed severaltaxonomic groups that are presently considered as separate families Edmundsrsquo workon mayflies was significant also in this area He proposed one of the first checklists ofmayflies from North America and Mexico (Edmunds and Allen 1957a) Hiscontribution to the biogeography and evolution of Ephemeroptera is still largelyaccurate (Edmunds 1972 1975) In 1976 he published in collaboration with Jensenand Berner a book with all the species known at that time from North and CentralAmerica (Edmunds et al 1976) In the Baetidae he described and reassigned severalspecies (Edmunds 1954 Edmunds and Allen 1957b)

In the Nearctic region contemporary authors such as Jacobus (Jacobus andMcCafferty 2005a 2005b 2006) Lugo-Ortiz and McCafferty (Lugo-Ortiz andMcCafferty 1998d) Waltz (Waltz and McCafferty 1986 1987b 1989) and Wiersema(Wiersema 2000 Wiersema and McCafferty 2004 Wiersema et al 2004) alsocontributed to the advanced knowledge of this fauna by the description of generaand species as well as several revisions with generic reassignments They sustain anactualised web page with a checklist and general information of the mayflies ofNorth and Central America (McCafferty 2009)

Figure 5 Specific and generic diversity in Nearctic realm Square total number of species(left scale) Triangle total number of genera (right scale)

50 J-L Gattolliat and C Nieto

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Sept

embe

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13

After 150 years of intensive works the systematics of Baetidae can be consideredas relatively well known even if some taxomomic problems are still not completelysolved The validity of some species needs to be reviewed a few of them have neverbeen mentioned since their original description

Australasian realm

The Australasian realm presently encompasses 40 known species all of themendemic except for Cloeon virens which is also distributed in the Oriental realmTwelve genera occur in this area only six of them being endemic (50) (Table 1)The non-endemic genera are also distributed in the Oriental region The presence ofsome typical Palearctic genera such as Centroptilum in Australasia is doubtful andshould be clarified in the near future

Offadens soror was the first species described from this region by Ulmer in 1908Following Ulmer Tillyard and Harker provided the only significant works onAustralian Baetidae (Tillyard 1936 Harker 1950 1954 1957) and until recently(Figure 6) Suter greatly improved the knowledge of the Ephemeroptera fauna in theAustralian region providing the first key for the larval stage of mayflies (Suter 1979)At that time the Baetidae encompassed only 12 species and five genera andinterestingly Bungona was the only genus which was endemic from this area all theother Australian species being attributed to Palearctic or Holarctic genera Later onSuter described new species and redescribed some genera (Suter 1986 2000 2001Suter and Pearson 2001) He also provided a key specifically for the larvae ofBaetidae which included several undescribed species and genera (Suter 1997)

As in other biogeographic regions Lugo-Ortiz and McCafferty played animportant role by describing several species and four endemic genera from thisregion (Lugo-Ortiz and McCafferty 1998e 1999a 1999c) and by reporting generasuch as Cloeodes or LabiobaetisPseudocloeon (Lugo-Ortiz and McCafferty 1998bLugo-Ortiz et al 1999) They also described several new taxa from New Guinea(Lugo-Ortiz and McCafferty 1999c Lugo-Ortiz et al 1999) Baetidae were one of thetwo families of mayflies reported from the Fiji Islands although it was only apreliminary study the diversity was relatively high with 12 species and three genera(Flowers 1990)

Figure 6 Specific and generic diversity in Australasian realm Square total number ofspecies (left scale) Triangle total number of genera (right scale)

Aquatic Insects 51

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13

Although the predaceous habit is very unusual in Baetidae two of the 12 generaof this region are carnivorous Echinobaetis from Sulawesi (Mol 1989) andMystaxyops from New Guinea (McCafferty and Sun 2005)

New Zealand is the only important landmass where mayflies are present andBaetidae have never been found This gap remains an enigma These islands are quiteisolated and possess endemic families such as Siphlaenigmatidae and Rallidentidaebut also the widespread family Leptophlebiidae The Baetidae were also consideredto be absent from New Caledonia (Peters 2001) However in material recentlycollected in standing waters half a dozen larvae of Cloeon were included(unpublished data) Because of the absence of Baetidae in previous intensivesamplings and the high power of dispersion of Cloeon it seems likely that a recentcolonisation of the island may be related to human activity If the Leptophlebiidaewere able to remain established on these islands after the break-off of the Gondwanaor to colonise the islands afterwards why were the Baetidae not able to do the sameThe question remains open

Despite recent improvements the fauna of this region remains insufficientlyknown in Australia and even more so on other islands and archipelagos ForAustralia a recent estimation established that only about half of the Australianspecies of Baetidae are described and probably some new genera still need to beestablished (Jeff Webb personal communication)

The biogeographic affinities of this region need to be analysed in detail All thenon-endemic genera are also distributed in the Oriental region they are mainlydiversified in the North East of Australasia and had probably dispersed step by stepthrough South East Asia into Australia where their distribution is limited to theNorth (Lugo-Ortiz and McCafferty 1999b) The genus Cloeodes clearly connects thisarea with the history of Gondwana and the continents placed in the SouthHemisphere Endemic genera such Edmundsiops Offadens and Bungona may berelated to South American taxa (Jeff Webb personal communication)

Conclusion

Since the establishment of the Baetidae in 1815 their knowledge has been hugelyincreased due to the efforts of many authors around the world In 1999 Lugo-Ortizand McCafferty provided the first worldwide synthesis of this family They estimatedthe diversity of this family at 650 species and 87 genera (Lugo-Ortiz and McCafferty1999b) Ten years later the number of species has increased by almost one third toreach 894 species During these 10 years 142 new species were described mainlyfrom South America Madagascar and West Africa It means that the increase of thetotal number of species is mainly due to the description of new species but Lugo-Ortiz and McCafferty had also underestimated the number of species of Baetidae byabout 100 species During these 10 years 17 genera were described and four weresynonymised

Baetidae are mostly diversified and possess a much higher rate of genericendemism in the Tropics and Southern Hemisphere than in the NorthernHemisphere (Barber-James et al 2008) The Afrotropics and Neotropics possessby far the most diversified faunas Despite common Gondwanian origins thesefaunas have almost no affinities The Palearctic fauna shares most genera with theOriental andor Afrotropical faunas as well as the Nearctic fauna Stepwisecolonisation could have occurred through the Palearctic region Central America

52 J-L Gattolliat and C Nieto

Dow

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ded

by [

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ity]

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957

14

Sept

embe

r 20

13

constitutes a bridge between North and South Americas and is under the influence ofboth areas (McCafferty 1998) The affinities of the Nearctic and Neotropical faunasare mainly due to the presence of North American taxa in Central America TheOriental fauna is still only partially known regions of biogeographic importance andwith a high potential diversity such as the Indian subcontinent and South East Asiaremain almost completely unknown Local studies clearly indicate that the diversityis quite high but the distribution of the taxa remains unknown The Oriental realm istherefore probably the most promising and challenging area Even if the globaldiversity is probably lower in Australasia the lack of knowledge of the fauna isobvious especially in the tropical part of this area such as New Guinea

The knowledge on systematics and faunistic composition are the first inevitablesteps to understand and preserve a natural system Most of the remaining unknownor poorly known faunas are under a direct human threat This underlines theurgency of this work before these faunas are definitively lost this is especially true inmost of the tropical rain forest areas where a primordial aquatic fauna is still presentFor areas with sufficient knowledge it is also the duty of the systematists to offer thenecessary tools for a correct identification of the taxa We can only encourageeveryone to make our knowledge accessible to others scientists especially by creatingkeys and checklists at different systematic levels

A comprehensive reconstruction of the phylogeny of the Baetidae is also highlynecessary Such a reconstruction should be based on modern tools and include bothmolecular and morphological characters It is the only way to solve contradictorysuprageneric relationships Such studies would allow a more accurate understandingof the relationships between the different areas in a global biogeographic approachThey would help to understand the historical influence of Gondwana and Laurasiaon the mayfly fauna

Acknowledgements

We want to express our appreciation to Michel Sartori (Museum of Zoology LausanneSwitzerland) and Helen Barber-James (Albany Museum Grahamstown South Africa) foruseful comments on this paper and great help in the conception of the Baetidae databaseAline Pasche (Museum of Zoology Lausanne Switzerland) offered technical supportFinancial support from the Argentine National Council of Scientific and TechnologicalResearch (CONICET) and ANPCYT (Pict 524-Pict 528) to which the junior author belongsare acknowledged The electronic database lsquolsquoEphemeroptera of the Worldrsquorsquo of Nikita Kluge(httpwwwinsectabiopuruzEph-sppContentshtm) was of great help for cross-checkingof our own database

References

Agnew JD (1961) lsquoNew Baetidae (Ephem) from South Africarsquo Novos Taxa Entomologicos25 3ndash18

Agnew JD (1963) lsquoNew South African records of imperfectly known Baetidae (Ephem)rsquoHydrobiologia 22 41ndash46

Bae YJ (1997) lsquoTaxonomy of Cloeon and Procloeon (Ephemeroptera Baetidae) in KorearsquoThe Korean Journal of Systematics Biology 13 303ndash314

Bae YJ and Park SY (1998) lsquoAlainites Baetis Labiobaetis and Nigrobaetis(Ephemeroptera Baetidae) in Korearsquo The Korean Journal of Systematics Biology 141ndash12

Banks N (1900) lsquoNew genera and species of Nearctic Neuropteroid Insectsrsquo Transactions ofthe American Entomological Society 26 239ndash259

Banks N (1914) lsquoNew neuropteroid insects native and exoticrsquo Proceedings of the Academy ofNatural Sciences of Philadelphia 66 608ndash632

Aquatic Insects 53

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

We can consider that the faunas of South West and East Africa as well as ofMadagascar are relatively well known In all of these regions there are mostcertainly several species and genera that remain unknown because of restricteddistribution or unusual habitat The fauna of the whole Atlantic side of Africa (fromCameroon to Angola) remains almost completely unknown This fauna probablyencompasses some taxa with a broad distribution like Centroptiloides bifasciata orOphelmatostoma camerunse but it certainly contains several new taxa The area ofthe Ruwenzori mountain range is also poorly known and probably possesses a highdegree of endemism Almost no report of Baetidae is available from NortheasternAfrica north to Kenya

Oriental realm

The first species of Baetidae from the Oriental realm were described from SriLanka and India (Hagen 1858 Eaton 1883ndash1888) (Figure 3) In his work on themayflies of the Sunda Islands Ulmer (1939) provided the first comprehensivereview of an Oriental fauna with complete and accurate descriptions of new speciesat both larval and imaginal stages Kimmins (1947) and Gillies (1949) bothcontributed to the knowledge of the Baetidae from India by respectively describingthree and 11 species from the imaginal stage Apart from these works the Orientalfauna remained very poorly known until the 1980s Muller-Liebenau describedabout 60 new species from the larval stage (about 50 of the species presentlyknown from this area) and six new genera (Muller-Liebenau 1980a 1980b 19811982a 1982b 1982c 1983 1984a 1984b 1985 Muller-Liebenau and Heard 1979Muller-Liebenau and Morihara 1982 Muller-Liebenau and Hubbard 1985) Shemainly described species from Sri Lanka Malaysia and the Philippines Her workconstituted the major contribution to the knowledge of Oriental fauna Waltz andMcCafferty helped to clarify the supra-specific classification by establishing newgenera and complexes of genera (Waltz and McCafferty 1987a 1987c Waltz et al1994)

The Baetidae from Taiwan were the subject of an extensive study allowing thedescription of 14 species (Kang et al 1994 Kang and Yang 1996) The fauna fromrunning waters can be considered as well known but the fauna from still and

Figure 3 Specific and generic diversity in Oriental realm Square total number of species(left scale) Triangle total number of genera (right scale)

Aquatic Insects 47

Dow

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ded

by [

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at 0

957

14

Sept

embe

r 20

13

standing water remained unknown The fauna of Hong Kong has recently beensubject to close attention with the description of five new species (Tong and Dudgeon2000 2003)

Although some islands like Sri Lanka Taiwan or Hong Kong were the subject ofimportant studies the majority of the Oriental realm remains virtually unknownFor example only provisional checklists of Baetidae at generic level are available forVietnam and Thailand India especially the southern part of this subcontinentprobably possesses a much more diversified fauna than the scattered reports seem toindicate There are also almost no reports from countries with potentially highdiversity such as Cambodia or Myanmar If we consider that in a 85 km2 area inKalimantan (Borneo) intensive sampling allowed the collection of 12 genera ofBaetidae five of them probably being new to science (Sartori et al 2003) we caneasily imagine that only a tiny part of Oriental fauna is presently known Mostfaunas are too poorly known to correctly estimate the distribution of the differentspecimens but preliminary results based on our own collections seem to indicate thatat least some species show a wide distribution including a great part of South EastAsia It is extremely difficult to estimate the specific and generic diversity of theOriental realm even for restricted regions it is probably not overstated to considerthat the species presently known represent less than a half or even a third of the realdiversity

Neotropical realm

With a total surface of 19 million of km2 the Neotropical realm is the fourthbiogeographical region The Baetidae there now encompass 210 species and 27genera (Table 1) The first species of Baetidae from the Neotropics were describedby Pictet (1843ndash1845) Eaton (1871 1883ndash1888) and Weyenbergh (1883)(Figure 4) As for the Afrotropics the descriptions were based on restrictedcollections of imagines deposited in European museums Navas was the mostprolific author unfortunately he described the same species several times underdifferent names he described 26 species of Callibaetis but only eight are presentlyconsidering as valid (Domınguez et al 2006) Most of the species described bythese first authors have been transferred to new genera or synonymised

Figure 4 Specific and generic diversity in Neotropical realm Square total number of species(left scale) Triangle total number of genera (right scale)

48 J-L Gattolliat and C Nieto

Dow

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ded

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r 20

13

Traver and Edmunds (1968) greatly improved the knowledge of the highlydiversified genus Camelobaetidius Mol (1986) established the only carnivorousgenus for this region Harpagobaetis Lugo-Ortiz and McCafferty (1995 1996a1998c 1999e) described 12 new genera from the Neotropics other generahave been recently established by Nieto (2003b) and McCafferty and Baumgardner(2003) Finally Domınguez et al (2006) published the first comprehensivereview of all South America species of Ephemeroptera with a chapter on thefamily Baetidae including a key illustrations distribution and ecology for eachtaxon

The isolation of this continent for millions of years is one of the mostimportant reasons why 23 of the Neotropical genera and more than 90 of thespecies are endemics from this realm Only seven genera are shared with theNearctic region with a generally higher specific richness in the Neotropics(Apobaetis Baetodes Callibaetis Camelobaetidius and Paracloeodes) only Amer-icabaetis and Baetis are more diversified in the Nearctic (McCafferty 1998) SeveralSouth American species were originally described in the genus Baetis andPseudocloeon but they were all subsequently transferred to other genera such asCamelobaetidius Cloeodes Fallceon and Moribaetis (Lugo-Ortiz and McCafferty1999d McCafferty 2000) Baetis magnus is the only true Baetis which is present inthe Neotropical realm its distribution encompasses a great part of CentralAmerica and the South of USA Baetis is present therefore in all the realmsFallceon certainly also has a Panamerican distribution the generic attribution ofF candidus from Taiwan is clearly a mistake and consequently Fallceon is notpresent in the Oriental realm Cloeodes is in fact the only genus present in SouthAmerica with a wide distribution as species attributed to this genus have beenreported from the whole Pantropical area Most genera possessing an almostworldwide distribution such as Cloeon or LabiobaetisPseudocloeon are onlymissing in the Neotropics this absence also greatly contributes to the originality ofthe Neotropical fauna

Several Neotropical taxa present peculiar adaptations Aturbina is the only genusof the Baetidae without turbinate eyes Callibaetis is one of the few viviparous generain the Ephemeroptera Chane is a spectacular genus with labium and maxillaeextremely well developed for collecting and filtering (Nieto 2003b) presentingsimilarities with the Afroptropical genus Ophelmatostoma (Gillies et al 1990) Mayo(1973) described the first species collected in a waterfall Baetis ellenae latertransferred to Mayobaetis (Waltz and McCafferty 1985) Cloeodes hydation istolerant to short periods of habitat desiccation this species occurs in smalltemporary rock pools (Nolte et al 1996)

Knowledge of the Baetidae from this region has been greatly improved in the lastfew decades new species and genera were described (Dominique et al 2002 Nieto2003a 2003b 2004a 2004b Thomas and Peru 2003) and other genera were reviewed(Nieto 2004c Nieto and Salles 2006 Nieto and Richard 2008) However there arestill regions or even countries with poorly known faunas such as Bolivia Chile PeruEcuador Uruguay and Venezuela these areas almost certainly possess someendemic taxa which remain to be found Recent samplings in Brazilian Amazoniaallowed the discovery of several new species (Salles and Batista 2004 Sallesand Francischetti 2004 Salles et al 2004 2005 Salles 2007 Salles and Polegatto2008) but a great part of this fauna remains unknown (Salles personalcommunication)

Aquatic Insects 49

Dow

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ded

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ate

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ity]

at 0

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Sept

embe

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13

Nearctic realm

Being the second largest biogeographic region (229 million km2) the Nearctic regionencompasses only 20 described genera (Table 1) The rate of endemism is also lowOnly five genera have a distribution restricted to this area (25) Seven genera areshared with the Neotropical region other genera present a distributionincluding the Palearctic andor Oriental areas Most genera are collector-gatherersor collector-scrapers no carnivorous Baetidae have been recorded from this regionFrom the 135 Nearctic species 114 are endemic (84) four Holarctic and 17Panamerican

The first ephemeropterologists working in this region were Hagen (1861) Walsh(1863a 1863b) and Eaton (1869 1870 1871 1881 1883ndash1888) Banks (1900 19141918) described eight species of Callibaetis five of which are still valid With 42species McDunnough was the main knowledgeable contributor to Nearctic Baetidae(McDunnough 1922 1923 1924 1925 1926 1929 1931 1932 1936 1939)(Figure 5)

The first accurate revision of all the species from North America was made byNeedham et al (1935) At that time the concept of Baetidae encompassed severaltaxonomic groups that are presently considered as separate families Edmundsrsquo workon mayflies was significant also in this area He proposed one of the first checklists ofmayflies from North America and Mexico (Edmunds and Allen 1957a) Hiscontribution to the biogeography and evolution of Ephemeroptera is still largelyaccurate (Edmunds 1972 1975) In 1976 he published in collaboration with Jensenand Berner a book with all the species known at that time from North and CentralAmerica (Edmunds et al 1976) In the Baetidae he described and reassigned severalspecies (Edmunds 1954 Edmunds and Allen 1957b)

In the Nearctic region contemporary authors such as Jacobus (Jacobus andMcCafferty 2005a 2005b 2006) Lugo-Ortiz and McCafferty (Lugo-Ortiz andMcCafferty 1998d) Waltz (Waltz and McCafferty 1986 1987b 1989) and Wiersema(Wiersema 2000 Wiersema and McCafferty 2004 Wiersema et al 2004) alsocontributed to the advanced knowledge of this fauna by the description of generaand species as well as several revisions with generic reassignments They sustain anactualised web page with a checklist and general information of the mayflies ofNorth and Central America (McCafferty 2009)

Figure 5 Specific and generic diversity in Nearctic realm Square total number of species(left scale) Triangle total number of genera (right scale)

50 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

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vers

ity]

at 0

957

14

Sept

embe

r 20

13

After 150 years of intensive works the systematics of Baetidae can be consideredas relatively well known even if some taxomomic problems are still not completelysolved The validity of some species needs to be reviewed a few of them have neverbeen mentioned since their original description

Australasian realm

The Australasian realm presently encompasses 40 known species all of themendemic except for Cloeon virens which is also distributed in the Oriental realmTwelve genera occur in this area only six of them being endemic (50) (Table 1)The non-endemic genera are also distributed in the Oriental region The presence ofsome typical Palearctic genera such as Centroptilum in Australasia is doubtful andshould be clarified in the near future

Offadens soror was the first species described from this region by Ulmer in 1908Following Ulmer Tillyard and Harker provided the only significant works onAustralian Baetidae (Tillyard 1936 Harker 1950 1954 1957) and until recently(Figure 6) Suter greatly improved the knowledge of the Ephemeroptera fauna in theAustralian region providing the first key for the larval stage of mayflies (Suter 1979)At that time the Baetidae encompassed only 12 species and five genera andinterestingly Bungona was the only genus which was endemic from this area all theother Australian species being attributed to Palearctic or Holarctic genera Later onSuter described new species and redescribed some genera (Suter 1986 2000 2001Suter and Pearson 2001) He also provided a key specifically for the larvae ofBaetidae which included several undescribed species and genera (Suter 1997)

As in other biogeographic regions Lugo-Ortiz and McCafferty played animportant role by describing several species and four endemic genera from thisregion (Lugo-Ortiz and McCafferty 1998e 1999a 1999c) and by reporting generasuch as Cloeodes or LabiobaetisPseudocloeon (Lugo-Ortiz and McCafferty 1998bLugo-Ortiz et al 1999) They also described several new taxa from New Guinea(Lugo-Ortiz and McCafferty 1999c Lugo-Ortiz et al 1999) Baetidae were one of thetwo families of mayflies reported from the Fiji Islands although it was only apreliminary study the diversity was relatively high with 12 species and three genera(Flowers 1990)

Figure 6 Specific and generic diversity in Australasian realm Square total number ofspecies (left scale) Triangle total number of genera (right scale)

Aquatic Insects 51

Dow

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ded

by [

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at 0

957

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Sept

embe

r 20

13

Although the predaceous habit is very unusual in Baetidae two of the 12 generaof this region are carnivorous Echinobaetis from Sulawesi (Mol 1989) andMystaxyops from New Guinea (McCafferty and Sun 2005)

New Zealand is the only important landmass where mayflies are present andBaetidae have never been found This gap remains an enigma These islands are quiteisolated and possess endemic families such as Siphlaenigmatidae and Rallidentidaebut also the widespread family Leptophlebiidae The Baetidae were also consideredto be absent from New Caledonia (Peters 2001) However in material recentlycollected in standing waters half a dozen larvae of Cloeon were included(unpublished data) Because of the absence of Baetidae in previous intensivesamplings and the high power of dispersion of Cloeon it seems likely that a recentcolonisation of the island may be related to human activity If the Leptophlebiidaewere able to remain established on these islands after the break-off of the Gondwanaor to colonise the islands afterwards why were the Baetidae not able to do the sameThe question remains open

Despite recent improvements the fauna of this region remains insufficientlyknown in Australia and even more so on other islands and archipelagos ForAustralia a recent estimation established that only about half of the Australianspecies of Baetidae are described and probably some new genera still need to beestablished (Jeff Webb personal communication)

The biogeographic affinities of this region need to be analysed in detail All thenon-endemic genera are also distributed in the Oriental region they are mainlydiversified in the North East of Australasia and had probably dispersed step by stepthrough South East Asia into Australia where their distribution is limited to theNorth (Lugo-Ortiz and McCafferty 1999b) The genus Cloeodes clearly connects thisarea with the history of Gondwana and the continents placed in the SouthHemisphere Endemic genera such Edmundsiops Offadens and Bungona may berelated to South American taxa (Jeff Webb personal communication)

Conclusion

Since the establishment of the Baetidae in 1815 their knowledge has been hugelyincreased due to the efforts of many authors around the world In 1999 Lugo-Ortizand McCafferty provided the first worldwide synthesis of this family They estimatedthe diversity of this family at 650 species and 87 genera (Lugo-Ortiz and McCafferty1999b) Ten years later the number of species has increased by almost one third toreach 894 species During these 10 years 142 new species were described mainlyfrom South America Madagascar and West Africa It means that the increase of thetotal number of species is mainly due to the description of new species but Lugo-Ortiz and McCafferty had also underestimated the number of species of Baetidae byabout 100 species During these 10 years 17 genera were described and four weresynonymised

Baetidae are mostly diversified and possess a much higher rate of genericendemism in the Tropics and Southern Hemisphere than in the NorthernHemisphere (Barber-James et al 2008) The Afrotropics and Neotropics possessby far the most diversified faunas Despite common Gondwanian origins thesefaunas have almost no affinities The Palearctic fauna shares most genera with theOriental andor Afrotropical faunas as well as the Nearctic fauna Stepwisecolonisation could have occurred through the Palearctic region Central America

52 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

constitutes a bridge between North and South Americas and is under the influence ofboth areas (McCafferty 1998) The affinities of the Nearctic and Neotropical faunasare mainly due to the presence of North American taxa in Central America TheOriental fauna is still only partially known regions of biogeographic importance andwith a high potential diversity such as the Indian subcontinent and South East Asiaremain almost completely unknown Local studies clearly indicate that the diversityis quite high but the distribution of the taxa remains unknown The Oriental realm istherefore probably the most promising and challenging area Even if the globaldiversity is probably lower in Australasia the lack of knowledge of the fauna isobvious especially in the tropical part of this area such as New Guinea

The knowledge on systematics and faunistic composition are the first inevitablesteps to understand and preserve a natural system Most of the remaining unknownor poorly known faunas are under a direct human threat This underlines theurgency of this work before these faunas are definitively lost this is especially true inmost of the tropical rain forest areas where a primordial aquatic fauna is still presentFor areas with sufficient knowledge it is also the duty of the systematists to offer thenecessary tools for a correct identification of the taxa We can only encourageeveryone to make our knowledge accessible to others scientists especially by creatingkeys and checklists at different systematic levels

A comprehensive reconstruction of the phylogeny of the Baetidae is also highlynecessary Such a reconstruction should be based on modern tools and include bothmolecular and morphological characters It is the only way to solve contradictorysuprageneric relationships Such studies would allow a more accurate understandingof the relationships between the different areas in a global biogeographic approachThey would help to understand the historical influence of Gondwana and Laurasiaon the mayfly fauna

Acknowledgements

We want to express our appreciation to Michel Sartori (Museum of Zoology LausanneSwitzerland) and Helen Barber-James (Albany Museum Grahamstown South Africa) foruseful comments on this paper and great help in the conception of the Baetidae databaseAline Pasche (Museum of Zoology Lausanne Switzerland) offered technical supportFinancial support from the Argentine National Council of Scientific and TechnologicalResearch (CONICET) and ANPCYT (Pict 524-Pict 528) to which the junior author belongsare acknowledged The electronic database lsquolsquoEphemeroptera of the Worldrsquorsquo of Nikita Kluge(httpwwwinsectabiopuruzEph-sppContentshtm) was of great help for cross-checkingof our own database

References

Agnew JD (1961) lsquoNew Baetidae (Ephem) from South Africarsquo Novos Taxa Entomologicos25 3ndash18

Agnew JD (1963) lsquoNew South African records of imperfectly known Baetidae (Ephem)rsquoHydrobiologia 22 41ndash46

Bae YJ (1997) lsquoTaxonomy of Cloeon and Procloeon (Ephemeroptera Baetidae) in KorearsquoThe Korean Journal of Systematics Biology 13 303ndash314

Bae YJ and Park SY (1998) lsquoAlainites Baetis Labiobaetis and Nigrobaetis(Ephemeroptera Baetidae) in Korearsquo The Korean Journal of Systematics Biology 141ndash12

Banks N (1900) lsquoNew genera and species of Nearctic Neuropteroid Insectsrsquo Transactions ofthe American Entomological Society 26 239ndash259

Banks N (1914) lsquoNew neuropteroid insects native and exoticrsquo Proceedings of the Academy ofNatural Sciences of Philadelphia 66 608ndash632

Aquatic Insects 53

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Sept

embe

r 20

13

standing water remained unknown The fauna of Hong Kong has recently beensubject to close attention with the description of five new species (Tong and Dudgeon2000 2003)

Although some islands like Sri Lanka Taiwan or Hong Kong were the subject ofimportant studies the majority of the Oriental realm remains virtually unknownFor example only provisional checklists of Baetidae at generic level are available forVietnam and Thailand India especially the southern part of this subcontinentprobably possesses a much more diversified fauna than the scattered reports seem toindicate There are also almost no reports from countries with potentially highdiversity such as Cambodia or Myanmar If we consider that in a 85 km2 area inKalimantan (Borneo) intensive sampling allowed the collection of 12 genera ofBaetidae five of them probably being new to science (Sartori et al 2003) we caneasily imagine that only a tiny part of Oriental fauna is presently known Mostfaunas are too poorly known to correctly estimate the distribution of the differentspecimens but preliminary results based on our own collections seem to indicate thatat least some species show a wide distribution including a great part of South EastAsia It is extremely difficult to estimate the specific and generic diversity of theOriental realm even for restricted regions it is probably not overstated to considerthat the species presently known represent less than a half or even a third of the realdiversity

Neotropical realm

With a total surface of 19 million of km2 the Neotropical realm is the fourthbiogeographical region The Baetidae there now encompass 210 species and 27genera (Table 1) The first species of Baetidae from the Neotropics were describedby Pictet (1843ndash1845) Eaton (1871 1883ndash1888) and Weyenbergh (1883)(Figure 4) As for the Afrotropics the descriptions were based on restrictedcollections of imagines deposited in European museums Navas was the mostprolific author unfortunately he described the same species several times underdifferent names he described 26 species of Callibaetis but only eight are presentlyconsidering as valid (Domınguez et al 2006) Most of the species described bythese first authors have been transferred to new genera or synonymised

Figure 4 Specific and generic diversity in Neotropical realm Square total number of species(left scale) Triangle total number of genera (right scale)

48 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Traver and Edmunds (1968) greatly improved the knowledge of the highlydiversified genus Camelobaetidius Mol (1986) established the only carnivorousgenus for this region Harpagobaetis Lugo-Ortiz and McCafferty (1995 1996a1998c 1999e) described 12 new genera from the Neotropics other generahave been recently established by Nieto (2003b) and McCafferty and Baumgardner(2003) Finally Domınguez et al (2006) published the first comprehensivereview of all South America species of Ephemeroptera with a chapter on thefamily Baetidae including a key illustrations distribution and ecology for eachtaxon

The isolation of this continent for millions of years is one of the mostimportant reasons why 23 of the Neotropical genera and more than 90 of thespecies are endemics from this realm Only seven genera are shared with theNearctic region with a generally higher specific richness in the Neotropics(Apobaetis Baetodes Callibaetis Camelobaetidius and Paracloeodes) only Amer-icabaetis and Baetis are more diversified in the Nearctic (McCafferty 1998) SeveralSouth American species were originally described in the genus Baetis andPseudocloeon but they were all subsequently transferred to other genera such asCamelobaetidius Cloeodes Fallceon and Moribaetis (Lugo-Ortiz and McCafferty1999d McCafferty 2000) Baetis magnus is the only true Baetis which is present inthe Neotropical realm its distribution encompasses a great part of CentralAmerica and the South of USA Baetis is present therefore in all the realmsFallceon certainly also has a Panamerican distribution the generic attribution ofF candidus from Taiwan is clearly a mistake and consequently Fallceon is notpresent in the Oriental realm Cloeodes is in fact the only genus present in SouthAmerica with a wide distribution as species attributed to this genus have beenreported from the whole Pantropical area Most genera possessing an almostworldwide distribution such as Cloeon or LabiobaetisPseudocloeon are onlymissing in the Neotropics this absence also greatly contributes to the originality ofthe Neotropical fauna

Several Neotropical taxa present peculiar adaptations Aturbina is the only genusof the Baetidae without turbinate eyes Callibaetis is one of the few viviparous generain the Ephemeroptera Chane is a spectacular genus with labium and maxillaeextremely well developed for collecting and filtering (Nieto 2003b) presentingsimilarities with the Afroptropical genus Ophelmatostoma (Gillies et al 1990) Mayo(1973) described the first species collected in a waterfall Baetis ellenae latertransferred to Mayobaetis (Waltz and McCafferty 1985) Cloeodes hydation istolerant to short periods of habitat desiccation this species occurs in smalltemporary rock pools (Nolte et al 1996)

Knowledge of the Baetidae from this region has been greatly improved in the lastfew decades new species and genera were described (Dominique et al 2002 Nieto2003a 2003b 2004a 2004b Thomas and Peru 2003) and other genera were reviewed(Nieto 2004c Nieto and Salles 2006 Nieto and Richard 2008) However there arestill regions or even countries with poorly known faunas such as Bolivia Chile PeruEcuador Uruguay and Venezuela these areas almost certainly possess someendemic taxa which remain to be found Recent samplings in Brazilian Amazoniaallowed the discovery of several new species (Salles and Batista 2004 Sallesand Francischetti 2004 Salles et al 2004 2005 Salles 2007 Salles and Polegatto2008) but a great part of this fauna remains unknown (Salles personalcommunication)

Aquatic Insects 49

Dow

nloa

ded

by [

Col

orad

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ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Nearctic realm

Being the second largest biogeographic region (229 million km2) the Nearctic regionencompasses only 20 described genera (Table 1) The rate of endemism is also lowOnly five genera have a distribution restricted to this area (25) Seven genera areshared with the Neotropical region other genera present a distributionincluding the Palearctic andor Oriental areas Most genera are collector-gatherersor collector-scrapers no carnivorous Baetidae have been recorded from this regionFrom the 135 Nearctic species 114 are endemic (84) four Holarctic and 17Panamerican

The first ephemeropterologists working in this region were Hagen (1861) Walsh(1863a 1863b) and Eaton (1869 1870 1871 1881 1883ndash1888) Banks (1900 19141918) described eight species of Callibaetis five of which are still valid With 42species McDunnough was the main knowledgeable contributor to Nearctic Baetidae(McDunnough 1922 1923 1924 1925 1926 1929 1931 1932 1936 1939)(Figure 5)

The first accurate revision of all the species from North America was made byNeedham et al (1935) At that time the concept of Baetidae encompassed severaltaxonomic groups that are presently considered as separate families Edmundsrsquo workon mayflies was significant also in this area He proposed one of the first checklists ofmayflies from North America and Mexico (Edmunds and Allen 1957a) Hiscontribution to the biogeography and evolution of Ephemeroptera is still largelyaccurate (Edmunds 1972 1975) In 1976 he published in collaboration with Jensenand Berner a book with all the species known at that time from North and CentralAmerica (Edmunds et al 1976) In the Baetidae he described and reassigned severalspecies (Edmunds 1954 Edmunds and Allen 1957b)

In the Nearctic region contemporary authors such as Jacobus (Jacobus andMcCafferty 2005a 2005b 2006) Lugo-Ortiz and McCafferty (Lugo-Ortiz andMcCafferty 1998d) Waltz (Waltz and McCafferty 1986 1987b 1989) and Wiersema(Wiersema 2000 Wiersema and McCafferty 2004 Wiersema et al 2004) alsocontributed to the advanced knowledge of this fauna by the description of generaand species as well as several revisions with generic reassignments They sustain anactualised web page with a checklist and general information of the mayflies ofNorth and Central America (McCafferty 2009)

Figure 5 Specific and generic diversity in Nearctic realm Square total number of species(left scale) Triangle total number of genera (right scale)

50 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

After 150 years of intensive works the systematics of Baetidae can be consideredas relatively well known even if some taxomomic problems are still not completelysolved The validity of some species needs to be reviewed a few of them have neverbeen mentioned since their original description

Australasian realm

The Australasian realm presently encompasses 40 known species all of themendemic except for Cloeon virens which is also distributed in the Oriental realmTwelve genera occur in this area only six of them being endemic (50) (Table 1)The non-endemic genera are also distributed in the Oriental region The presence ofsome typical Palearctic genera such as Centroptilum in Australasia is doubtful andshould be clarified in the near future

Offadens soror was the first species described from this region by Ulmer in 1908Following Ulmer Tillyard and Harker provided the only significant works onAustralian Baetidae (Tillyard 1936 Harker 1950 1954 1957) and until recently(Figure 6) Suter greatly improved the knowledge of the Ephemeroptera fauna in theAustralian region providing the first key for the larval stage of mayflies (Suter 1979)At that time the Baetidae encompassed only 12 species and five genera andinterestingly Bungona was the only genus which was endemic from this area all theother Australian species being attributed to Palearctic or Holarctic genera Later onSuter described new species and redescribed some genera (Suter 1986 2000 2001Suter and Pearson 2001) He also provided a key specifically for the larvae ofBaetidae which included several undescribed species and genera (Suter 1997)

As in other biogeographic regions Lugo-Ortiz and McCafferty played animportant role by describing several species and four endemic genera from thisregion (Lugo-Ortiz and McCafferty 1998e 1999a 1999c) and by reporting generasuch as Cloeodes or LabiobaetisPseudocloeon (Lugo-Ortiz and McCafferty 1998bLugo-Ortiz et al 1999) They also described several new taxa from New Guinea(Lugo-Ortiz and McCafferty 1999c Lugo-Ortiz et al 1999) Baetidae were one of thetwo families of mayflies reported from the Fiji Islands although it was only apreliminary study the diversity was relatively high with 12 species and three genera(Flowers 1990)

Figure 6 Specific and generic diversity in Australasian realm Square total number ofspecies (left scale) Triangle total number of genera (right scale)

Aquatic Insects 51

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Although the predaceous habit is very unusual in Baetidae two of the 12 generaof this region are carnivorous Echinobaetis from Sulawesi (Mol 1989) andMystaxyops from New Guinea (McCafferty and Sun 2005)

New Zealand is the only important landmass where mayflies are present andBaetidae have never been found This gap remains an enigma These islands are quiteisolated and possess endemic families such as Siphlaenigmatidae and Rallidentidaebut also the widespread family Leptophlebiidae The Baetidae were also consideredto be absent from New Caledonia (Peters 2001) However in material recentlycollected in standing waters half a dozen larvae of Cloeon were included(unpublished data) Because of the absence of Baetidae in previous intensivesamplings and the high power of dispersion of Cloeon it seems likely that a recentcolonisation of the island may be related to human activity If the Leptophlebiidaewere able to remain established on these islands after the break-off of the Gondwanaor to colonise the islands afterwards why were the Baetidae not able to do the sameThe question remains open

Despite recent improvements the fauna of this region remains insufficientlyknown in Australia and even more so on other islands and archipelagos ForAustralia a recent estimation established that only about half of the Australianspecies of Baetidae are described and probably some new genera still need to beestablished (Jeff Webb personal communication)

The biogeographic affinities of this region need to be analysed in detail All thenon-endemic genera are also distributed in the Oriental region they are mainlydiversified in the North East of Australasia and had probably dispersed step by stepthrough South East Asia into Australia where their distribution is limited to theNorth (Lugo-Ortiz and McCafferty 1999b) The genus Cloeodes clearly connects thisarea with the history of Gondwana and the continents placed in the SouthHemisphere Endemic genera such Edmundsiops Offadens and Bungona may berelated to South American taxa (Jeff Webb personal communication)

Conclusion

Since the establishment of the Baetidae in 1815 their knowledge has been hugelyincreased due to the efforts of many authors around the world In 1999 Lugo-Ortizand McCafferty provided the first worldwide synthesis of this family They estimatedthe diversity of this family at 650 species and 87 genera (Lugo-Ortiz and McCafferty1999b) Ten years later the number of species has increased by almost one third toreach 894 species During these 10 years 142 new species were described mainlyfrom South America Madagascar and West Africa It means that the increase of thetotal number of species is mainly due to the description of new species but Lugo-Ortiz and McCafferty had also underestimated the number of species of Baetidae byabout 100 species During these 10 years 17 genera were described and four weresynonymised

Baetidae are mostly diversified and possess a much higher rate of genericendemism in the Tropics and Southern Hemisphere than in the NorthernHemisphere (Barber-James et al 2008) The Afrotropics and Neotropics possessby far the most diversified faunas Despite common Gondwanian origins thesefaunas have almost no affinities The Palearctic fauna shares most genera with theOriental andor Afrotropical faunas as well as the Nearctic fauna Stepwisecolonisation could have occurred through the Palearctic region Central America

52 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

constitutes a bridge between North and South Americas and is under the influence ofboth areas (McCafferty 1998) The affinities of the Nearctic and Neotropical faunasare mainly due to the presence of North American taxa in Central America TheOriental fauna is still only partially known regions of biogeographic importance andwith a high potential diversity such as the Indian subcontinent and South East Asiaremain almost completely unknown Local studies clearly indicate that the diversityis quite high but the distribution of the taxa remains unknown The Oriental realm istherefore probably the most promising and challenging area Even if the globaldiversity is probably lower in Australasia the lack of knowledge of the fauna isobvious especially in the tropical part of this area such as New Guinea

The knowledge on systematics and faunistic composition are the first inevitablesteps to understand and preserve a natural system Most of the remaining unknownor poorly known faunas are under a direct human threat This underlines theurgency of this work before these faunas are definitively lost this is especially true inmost of the tropical rain forest areas where a primordial aquatic fauna is still presentFor areas with sufficient knowledge it is also the duty of the systematists to offer thenecessary tools for a correct identification of the taxa We can only encourageeveryone to make our knowledge accessible to others scientists especially by creatingkeys and checklists at different systematic levels

A comprehensive reconstruction of the phylogeny of the Baetidae is also highlynecessary Such a reconstruction should be based on modern tools and include bothmolecular and morphological characters It is the only way to solve contradictorysuprageneric relationships Such studies would allow a more accurate understandingof the relationships between the different areas in a global biogeographic approachThey would help to understand the historical influence of Gondwana and Laurasiaon the mayfly fauna

Acknowledgements

We want to express our appreciation to Michel Sartori (Museum of Zoology LausanneSwitzerland) and Helen Barber-James (Albany Museum Grahamstown South Africa) foruseful comments on this paper and great help in the conception of the Baetidae databaseAline Pasche (Museum of Zoology Lausanne Switzerland) offered technical supportFinancial support from the Argentine National Council of Scientific and TechnologicalResearch (CONICET) and ANPCYT (Pict 524-Pict 528) to which the junior author belongsare acknowledged The electronic database lsquolsquoEphemeroptera of the Worldrsquorsquo of Nikita Kluge(httpwwwinsectabiopuruzEph-sppContentshtm) was of great help for cross-checkingof our own database

References

Agnew JD (1961) lsquoNew Baetidae (Ephem) from South Africarsquo Novos Taxa Entomologicos25 3ndash18

Agnew JD (1963) lsquoNew South African records of imperfectly known Baetidae (Ephem)rsquoHydrobiologia 22 41ndash46

Bae YJ (1997) lsquoTaxonomy of Cloeon and Procloeon (Ephemeroptera Baetidae) in KorearsquoThe Korean Journal of Systematics Biology 13 303ndash314

Bae YJ and Park SY (1998) lsquoAlainites Baetis Labiobaetis and Nigrobaetis(Ephemeroptera Baetidae) in Korearsquo The Korean Journal of Systematics Biology 141ndash12

Banks N (1900) lsquoNew genera and species of Nearctic Neuropteroid Insectsrsquo Transactions ofthe American Entomological Society 26 239ndash259

Banks N (1914) lsquoNew neuropteroid insects native and exoticrsquo Proceedings of the Academy ofNatural Sciences of Philadelphia 66 608ndash632

Aquatic Insects 53

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Traver and Edmunds (1968) greatly improved the knowledge of the highlydiversified genus Camelobaetidius Mol (1986) established the only carnivorousgenus for this region Harpagobaetis Lugo-Ortiz and McCafferty (1995 1996a1998c 1999e) described 12 new genera from the Neotropics other generahave been recently established by Nieto (2003b) and McCafferty and Baumgardner(2003) Finally Domınguez et al (2006) published the first comprehensivereview of all South America species of Ephemeroptera with a chapter on thefamily Baetidae including a key illustrations distribution and ecology for eachtaxon

The isolation of this continent for millions of years is one of the mostimportant reasons why 23 of the Neotropical genera and more than 90 of thespecies are endemics from this realm Only seven genera are shared with theNearctic region with a generally higher specific richness in the Neotropics(Apobaetis Baetodes Callibaetis Camelobaetidius and Paracloeodes) only Amer-icabaetis and Baetis are more diversified in the Nearctic (McCafferty 1998) SeveralSouth American species were originally described in the genus Baetis andPseudocloeon but they were all subsequently transferred to other genera such asCamelobaetidius Cloeodes Fallceon and Moribaetis (Lugo-Ortiz and McCafferty1999d McCafferty 2000) Baetis magnus is the only true Baetis which is present inthe Neotropical realm its distribution encompasses a great part of CentralAmerica and the South of USA Baetis is present therefore in all the realmsFallceon certainly also has a Panamerican distribution the generic attribution ofF candidus from Taiwan is clearly a mistake and consequently Fallceon is notpresent in the Oriental realm Cloeodes is in fact the only genus present in SouthAmerica with a wide distribution as species attributed to this genus have beenreported from the whole Pantropical area Most genera possessing an almostworldwide distribution such as Cloeon or LabiobaetisPseudocloeon are onlymissing in the Neotropics this absence also greatly contributes to the originality ofthe Neotropical fauna

Several Neotropical taxa present peculiar adaptations Aturbina is the only genusof the Baetidae without turbinate eyes Callibaetis is one of the few viviparous generain the Ephemeroptera Chane is a spectacular genus with labium and maxillaeextremely well developed for collecting and filtering (Nieto 2003b) presentingsimilarities with the Afroptropical genus Ophelmatostoma (Gillies et al 1990) Mayo(1973) described the first species collected in a waterfall Baetis ellenae latertransferred to Mayobaetis (Waltz and McCafferty 1985) Cloeodes hydation istolerant to short periods of habitat desiccation this species occurs in smalltemporary rock pools (Nolte et al 1996)

Knowledge of the Baetidae from this region has been greatly improved in the lastfew decades new species and genera were described (Dominique et al 2002 Nieto2003a 2003b 2004a 2004b Thomas and Peru 2003) and other genera were reviewed(Nieto 2004c Nieto and Salles 2006 Nieto and Richard 2008) However there arestill regions or even countries with poorly known faunas such as Bolivia Chile PeruEcuador Uruguay and Venezuela these areas almost certainly possess someendemic taxa which remain to be found Recent samplings in Brazilian Amazoniaallowed the discovery of several new species (Salles and Batista 2004 Sallesand Francischetti 2004 Salles et al 2004 2005 Salles 2007 Salles and Polegatto2008) but a great part of this fauna remains unknown (Salles personalcommunication)

Aquatic Insects 49

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Nearctic realm

Being the second largest biogeographic region (229 million km2) the Nearctic regionencompasses only 20 described genera (Table 1) The rate of endemism is also lowOnly five genera have a distribution restricted to this area (25) Seven genera areshared with the Neotropical region other genera present a distributionincluding the Palearctic andor Oriental areas Most genera are collector-gatherersor collector-scrapers no carnivorous Baetidae have been recorded from this regionFrom the 135 Nearctic species 114 are endemic (84) four Holarctic and 17Panamerican

The first ephemeropterologists working in this region were Hagen (1861) Walsh(1863a 1863b) and Eaton (1869 1870 1871 1881 1883ndash1888) Banks (1900 19141918) described eight species of Callibaetis five of which are still valid With 42species McDunnough was the main knowledgeable contributor to Nearctic Baetidae(McDunnough 1922 1923 1924 1925 1926 1929 1931 1932 1936 1939)(Figure 5)

The first accurate revision of all the species from North America was made byNeedham et al (1935) At that time the concept of Baetidae encompassed severaltaxonomic groups that are presently considered as separate families Edmundsrsquo workon mayflies was significant also in this area He proposed one of the first checklists ofmayflies from North America and Mexico (Edmunds and Allen 1957a) Hiscontribution to the biogeography and evolution of Ephemeroptera is still largelyaccurate (Edmunds 1972 1975) In 1976 he published in collaboration with Jensenand Berner a book with all the species known at that time from North and CentralAmerica (Edmunds et al 1976) In the Baetidae he described and reassigned severalspecies (Edmunds 1954 Edmunds and Allen 1957b)

In the Nearctic region contemporary authors such as Jacobus (Jacobus andMcCafferty 2005a 2005b 2006) Lugo-Ortiz and McCafferty (Lugo-Ortiz andMcCafferty 1998d) Waltz (Waltz and McCafferty 1986 1987b 1989) and Wiersema(Wiersema 2000 Wiersema and McCafferty 2004 Wiersema et al 2004) alsocontributed to the advanced knowledge of this fauna by the description of generaand species as well as several revisions with generic reassignments They sustain anactualised web page with a checklist and general information of the mayflies ofNorth and Central America (McCafferty 2009)

Figure 5 Specific and generic diversity in Nearctic realm Square total number of species(left scale) Triangle total number of genera (right scale)

50 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

After 150 years of intensive works the systematics of Baetidae can be consideredas relatively well known even if some taxomomic problems are still not completelysolved The validity of some species needs to be reviewed a few of them have neverbeen mentioned since their original description

Australasian realm

The Australasian realm presently encompasses 40 known species all of themendemic except for Cloeon virens which is also distributed in the Oriental realmTwelve genera occur in this area only six of them being endemic (50) (Table 1)The non-endemic genera are also distributed in the Oriental region The presence ofsome typical Palearctic genera such as Centroptilum in Australasia is doubtful andshould be clarified in the near future

Offadens soror was the first species described from this region by Ulmer in 1908Following Ulmer Tillyard and Harker provided the only significant works onAustralian Baetidae (Tillyard 1936 Harker 1950 1954 1957) and until recently(Figure 6) Suter greatly improved the knowledge of the Ephemeroptera fauna in theAustralian region providing the first key for the larval stage of mayflies (Suter 1979)At that time the Baetidae encompassed only 12 species and five genera andinterestingly Bungona was the only genus which was endemic from this area all theother Australian species being attributed to Palearctic or Holarctic genera Later onSuter described new species and redescribed some genera (Suter 1986 2000 2001Suter and Pearson 2001) He also provided a key specifically for the larvae ofBaetidae which included several undescribed species and genera (Suter 1997)

As in other biogeographic regions Lugo-Ortiz and McCafferty played animportant role by describing several species and four endemic genera from thisregion (Lugo-Ortiz and McCafferty 1998e 1999a 1999c) and by reporting generasuch as Cloeodes or LabiobaetisPseudocloeon (Lugo-Ortiz and McCafferty 1998bLugo-Ortiz et al 1999) They also described several new taxa from New Guinea(Lugo-Ortiz and McCafferty 1999c Lugo-Ortiz et al 1999) Baetidae were one of thetwo families of mayflies reported from the Fiji Islands although it was only apreliminary study the diversity was relatively high with 12 species and three genera(Flowers 1990)

Figure 6 Specific and generic diversity in Australasian realm Square total number ofspecies (left scale) Triangle total number of genera (right scale)

Aquatic Insects 51

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Although the predaceous habit is very unusual in Baetidae two of the 12 generaof this region are carnivorous Echinobaetis from Sulawesi (Mol 1989) andMystaxyops from New Guinea (McCafferty and Sun 2005)

New Zealand is the only important landmass where mayflies are present andBaetidae have never been found This gap remains an enigma These islands are quiteisolated and possess endemic families such as Siphlaenigmatidae and Rallidentidaebut also the widespread family Leptophlebiidae The Baetidae were also consideredto be absent from New Caledonia (Peters 2001) However in material recentlycollected in standing waters half a dozen larvae of Cloeon were included(unpublished data) Because of the absence of Baetidae in previous intensivesamplings and the high power of dispersion of Cloeon it seems likely that a recentcolonisation of the island may be related to human activity If the Leptophlebiidaewere able to remain established on these islands after the break-off of the Gondwanaor to colonise the islands afterwards why were the Baetidae not able to do the sameThe question remains open

Despite recent improvements the fauna of this region remains insufficientlyknown in Australia and even more so on other islands and archipelagos ForAustralia a recent estimation established that only about half of the Australianspecies of Baetidae are described and probably some new genera still need to beestablished (Jeff Webb personal communication)

The biogeographic affinities of this region need to be analysed in detail All thenon-endemic genera are also distributed in the Oriental region they are mainlydiversified in the North East of Australasia and had probably dispersed step by stepthrough South East Asia into Australia where their distribution is limited to theNorth (Lugo-Ortiz and McCafferty 1999b) The genus Cloeodes clearly connects thisarea with the history of Gondwana and the continents placed in the SouthHemisphere Endemic genera such Edmundsiops Offadens and Bungona may berelated to South American taxa (Jeff Webb personal communication)

Conclusion

Since the establishment of the Baetidae in 1815 their knowledge has been hugelyincreased due to the efforts of many authors around the world In 1999 Lugo-Ortizand McCafferty provided the first worldwide synthesis of this family They estimatedthe diversity of this family at 650 species and 87 genera (Lugo-Ortiz and McCafferty1999b) Ten years later the number of species has increased by almost one third toreach 894 species During these 10 years 142 new species were described mainlyfrom South America Madagascar and West Africa It means that the increase of thetotal number of species is mainly due to the description of new species but Lugo-Ortiz and McCafferty had also underestimated the number of species of Baetidae byabout 100 species During these 10 years 17 genera were described and four weresynonymised

Baetidae are mostly diversified and possess a much higher rate of genericendemism in the Tropics and Southern Hemisphere than in the NorthernHemisphere (Barber-James et al 2008) The Afrotropics and Neotropics possessby far the most diversified faunas Despite common Gondwanian origins thesefaunas have almost no affinities The Palearctic fauna shares most genera with theOriental andor Afrotropical faunas as well as the Nearctic fauna Stepwisecolonisation could have occurred through the Palearctic region Central America

52 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

constitutes a bridge between North and South Americas and is under the influence ofboth areas (McCafferty 1998) The affinities of the Nearctic and Neotropical faunasare mainly due to the presence of North American taxa in Central America TheOriental fauna is still only partially known regions of biogeographic importance andwith a high potential diversity such as the Indian subcontinent and South East Asiaremain almost completely unknown Local studies clearly indicate that the diversityis quite high but the distribution of the taxa remains unknown The Oriental realm istherefore probably the most promising and challenging area Even if the globaldiversity is probably lower in Australasia the lack of knowledge of the fauna isobvious especially in the tropical part of this area such as New Guinea

The knowledge on systematics and faunistic composition are the first inevitablesteps to understand and preserve a natural system Most of the remaining unknownor poorly known faunas are under a direct human threat This underlines theurgency of this work before these faunas are definitively lost this is especially true inmost of the tropical rain forest areas where a primordial aquatic fauna is still presentFor areas with sufficient knowledge it is also the duty of the systematists to offer thenecessary tools for a correct identification of the taxa We can only encourageeveryone to make our knowledge accessible to others scientists especially by creatingkeys and checklists at different systematic levels

A comprehensive reconstruction of the phylogeny of the Baetidae is also highlynecessary Such a reconstruction should be based on modern tools and include bothmolecular and morphological characters It is the only way to solve contradictorysuprageneric relationships Such studies would allow a more accurate understandingof the relationships between the different areas in a global biogeographic approachThey would help to understand the historical influence of Gondwana and Laurasiaon the mayfly fauna

Acknowledgements

We want to express our appreciation to Michel Sartori (Museum of Zoology LausanneSwitzerland) and Helen Barber-James (Albany Museum Grahamstown South Africa) foruseful comments on this paper and great help in the conception of the Baetidae databaseAline Pasche (Museum of Zoology Lausanne Switzerland) offered technical supportFinancial support from the Argentine National Council of Scientific and TechnologicalResearch (CONICET) and ANPCYT (Pict 524-Pict 528) to which the junior author belongsare acknowledged The electronic database lsquolsquoEphemeroptera of the Worldrsquorsquo of Nikita Kluge(httpwwwinsectabiopuruzEph-sppContentshtm) was of great help for cross-checkingof our own database

References

Agnew JD (1961) lsquoNew Baetidae (Ephem) from South Africarsquo Novos Taxa Entomologicos25 3ndash18

Agnew JD (1963) lsquoNew South African records of imperfectly known Baetidae (Ephem)rsquoHydrobiologia 22 41ndash46

Bae YJ (1997) lsquoTaxonomy of Cloeon and Procloeon (Ephemeroptera Baetidae) in KorearsquoThe Korean Journal of Systematics Biology 13 303ndash314

Bae YJ and Park SY (1998) lsquoAlainites Baetis Labiobaetis and Nigrobaetis(Ephemeroptera Baetidae) in Korearsquo The Korean Journal of Systematics Biology 141ndash12

Banks N (1900) lsquoNew genera and species of Nearctic Neuropteroid Insectsrsquo Transactions ofthe American Entomological Society 26 239ndash259

Banks N (1914) lsquoNew neuropteroid insects native and exoticrsquo Proceedings of the Academy ofNatural Sciences of Philadelphia 66 608ndash632

Aquatic Insects 53

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Nearctic realm

Being the second largest biogeographic region (229 million km2) the Nearctic regionencompasses only 20 described genera (Table 1) The rate of endemism is also lowOnly five genera have a distribution restricted to this area (25) Seven genera areshared with the Neotropical region other genera present a distributionincluding the Palearctic andor Oriental areas Most genera are collector-gatherersor collector-scrapers no carnivorous Baetidae have been recorded from this regionFrom the 135 Nearctic species 114 are endemic (84) four Holarctic and 17Panamerican

The first ephemeropterologists working in this region were Hagen (1861) Walsh(1863a 1863b) and Eaton (1869 1870 1871 1881 1883ndash1888) Banks (1900 19141918) described eight species of Callibaetis five of which are still valid With 42species McDunnough was the main knowledgeable contributor to Nearctic Baetidae(McDunnough 1922 1923 1924 1925 1926 1929 1931 1932 1936 1939)(Figure 5)

The first accurate revision of all the species from North America was made byNeedham et al (1935) At that time the concept of Baetidae encompassed severaltaxonomic groups that are presently considered as separate families Edmundsrsquo workon mayflies was significant also in this area He proposed one of the first checklists ofmayflies from North America and Mexico (Edmunds and Allen 1957a) Hiscontribution to the biogeography and evolution of Ephemeroptera is still largelyaccurate (Edmunds 1972 1975) In 1976 he published in collaboration with Jensenand Berner a book with all the species known at that time from North and CentralAmerica (Edmunds et al 1976) In the Baetidae he described and reassigned severalspecies (Edmunds 1954 Edmunds and Allen 1957b)

In the Nearctic region contemporary authors such as Jacobus (Jacobus andMcCafferty 2005a 2005b 2006) Lugo-Ortiz and McCafferty (Lugo-Ortiz andMcCafferty 1998d) Waltz (Waltz and McCafferty 1986 1987b 1989) and Wiersema(Wiersema 2000 Wiersema and McCafferty 2004 Wiersema et al 2004) alsocontributed to the advanced knowledge of this fauna by the description of generaand species as well as several revisions with generic reassignments They sustain anactualised web page with a checklist and general information of the mayflies ofNorth and Central America (McCafferty 2009)

Figure 5 Specific and generic diversity in Nearctic realm Square total number of species(left scale) Triangle total number of genera (right scale)

50 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

After 150 years of intensive works the systematics of Baetidae can be consideredas relatively well known even if some taxomomic problems are still not completelysolved The validity of some species needs to be reviewed a few of them have neverbeen mentioned since their original description

Australasian realm

The Australasian realm presently encompasses 40 known species all of themendemic except for Cloeon virens which is also distributed in the Oriental realmTwelve genera occur in this area only six of them being endemic (50) (Table 1)The non-endemic genera are also distributed in the Oriental region The presence ofsome typical Palearctic genera such as Centroptilum in Australasia is doubtful andshould be clarified in the near future

Offadens soror was the first species described from this region by Ulmer in 1908Following Ulmer Tillyard and Harker provided the only significant works onAustralian Baetidae (Tillyard 1936 Harker 1950 1954 1957) and until recently(Figure 6) Suter greatly improved the knowledge of the Ephemeroptera fauna in theAustralian region providing the first key for the larval stage of mayflies (Suter 1979)At that time the Baetidae encompassed only 12 species and five genera andinterestingly Bungona was the only genus which was endemic from this area all theother Australian species being attributed to Palearctic or Holarctic genera Later onSuter described new species and redescribed some genera (Suter 1986 2000 2001Suter and Pearson 2001) He also provided a key specifically for the larvae ofBaetidae which included several undescribed species and genera (Suter 1997)

As in other biogeographic regions Lugo-Ortiz and McCafferty played animportant role by describing several species and four endemic genera from thisregion (Lugo-Ortiz and McCafferty 1998e 1999a 1999c) and by reporting generasuch as Cloeodes or LabiobaetisPseudocloeon (Lugo-Ortiz and McCafferty 1998bLugo-Ortiz et al 1999) They also described several new taxa from New Guinea(Lugo-Ortiz and McCafferty 1999c Lugo-Ortiz et al 1999) Baetidae were one of thetwo families of mayflies reported from the Fiji Islands although it was only apreliminary study the diversity was relatively high with 12 species and three genera(Flowers 1990)

Figure 6 Specific and generic diversity in Australasian realm Square total number ofspecies (left scale) Triangle total number of genera (right scale)

Aquatic Insects 51

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Although the predaceous habit is very unusual in Baetidae two of the 12 generaof this region are carnivorous Echinobaetis from Sulawesi (Mol 1989) andMystaxyops from New Guinea (McCafferty and Sun 2005)

New Zealand is the only important landmass where mayflies are present andBaetidae have never been found This gap remains an enigma These islands are quiteisolated and possess endemic families such as Siphlaenigmatidae and Rallidentidaebut also the widespread family Leptophlebiidae The Baetidae were also consideredto be absent from New Caledonia (Peters 2001) However in material recentlycollected in standing waters half a dozen larvae of Cloeon were included(unpublished data) Because of the absence of Baetidae in previous intensivesamplings and the high power of dispersion of Cloeon it seems likely that a recentcolonisation of the island may be related to human activity If the Leptophlebiidaewere able to remain established on these islands after the break-off of the Gondwanaor to colonise the islands afterwards why were the Baetidae not able to do the sameThe question remains open

Despite recent improvements the fauna of this region remains insufficientlyknown in Australia and even more so on other islands and archipelagos ForAustralia a recent estimation established that only about half of the Australianspecies of Baetidae are described and probably some new genera still need to beestablished (Jeff Webb personal communication)

The biogeographic affinities of this region need to be analysed in detail All thenon-endemic genera are also distributed in the Oriental region they are mainlydiversified in the North East of Australasia and had probably dispersed step by stepthrough South East Asia into Australia where their distribution is limited to theNorth (Lugo-Ortiz and McCafferty 1999b) The genus Cloeodes clearly connects thisarea with the history of Gondwana and the continents placed in the SouthHemisphere Endemic genera such Edmundsiops Offadens and Bungona may berelated to South American taxa (Jeff Webb personal communication)

Conclusion

Since the establishment of the Baetidae in 1815 their knowledge has been hugelyincreased due to the efforts of many authors around the world In 1999 Lugo-Ortizand McCafferty provided the first worldwide synthesis of this family They estimatedthe diversity of this family at 650 species and 87 genera (Lugo-Ortiz and McCafferty1999b) Ten years later the number of species has increased by almost one third toreach 894 species During these 10 years 142 new species were described mainlyfrom South America Madagascar and West Africa It means that the increase of thetotal number of species is mainly due to the description of new species but Lugo-Ortiz and McCafferty had also underestimated the number of species of Baetidae byabout 100 species During these 10 years 17 genera were described and four weresynonymised

Baetidae are mostly diversified and possess a much higher rate of genericendemism in the Tropics and Southern Hemisphere than in the NorthernHemisphere (Barber-James et al 2008) The Afrotropics and Neotropics possessby far the most diversified faunas Despite common Gondwanian origins thesefaunas have almost no affinities The Palearctic fauna shares most genera with theOriental andor Afrotropical faunas as well as the Nearctic fauna Stepwisecolonisation could have occurred through the Palearctic region Central America

52 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

constitutes a bridge between North and South Americas and is under the influence ofboth areas (McCafferty 1998) The affinities of the Nearctic and Neotropical faunasare mainly due to the presence of North American taxa in Central America TheOriental fauna is still only partially known regions of biogeographic importance andwith a high potential diversity such as the Indian subcontinent and South East Asiaremain almost completely unknown Local studies clearly indicate that the diversityis quite high but the distribution of the taxa remains unknown The Oriental realm istherefore probably the most promising and challenging area Even if the globaldiversity is probably lower in Australasia the lack of knowledge of the fauna isobvious especially in the tropical part of this area such as New Guinea

The knowledge on systematics and faunistic composition are the first inevitablesteps to understand and preserve a natural system Most of the remaining unknownor poorly known faunas are under a direct human threat This underlines theurgency of this work before these faunas are definitively lost this is especially true inmost of the tropical rain forest areas where a primordial aquatic fauna is still presentFor areas with sufficient knowledge it is also the duty of the systematists to offer thenecessary tools for a correct identification of the taxa We can only encourageeveryone to make our knowledge accessible to others scientists especially by creatingkeys and checklists at different systematic levels

A comprehensive reconstruction of the phylogeny of the Baetidae is also highlynecessary Such a reconstruction should be based on modern tools and include bothmolecular and morphological characters It is the only way to solve contradictorysuprageneric relationships Such studies would allow a more accurate understandingof the relationships between the different areas in a global biogeographic approachThey would help to understand the historical influence of Gondwana and Laurasiaon the mayfly fauna

Acknowledgements

We want to express our appreciation to Michel Sartori (Museum of Zoology LausanneSwitzerland) and Helen Barber-James (Albany Museum Grahamstown South Africa) foruseful comments on this paper and great help in the conception of the Baetidae databaseAline Pasche (Museum of Zoology Lausanne Switzerland) offered technical supportFinancial support from the Argentine National Council of Scientific and TechnologicalResearch (CONICET) and ANPCYT (Pict 524-Pict 528) to which the junior author belongsare acknowledged The electronic database lsquolsquoEphemeroptera of the Worldrsquorsquo of Nikita Kluge(httpwwwinsectabiopuruzEph-sppContentshtm) was of great help for cross-checkingof our own database

References

Agnew JD (1961) lsquoNew Baetidae (Ephem) from South Africarsquo Novos Taxa Entomologicos25 3ndash18

Agnew JD (1963) lsquoNew South African records of imperfectly known Baetidae (Ephem)rsquoHydrobiologia 22 41ndash46

Bae YJ (1997) lsquoTaxonomy of Cloeon and Procloeon (Ephemeroptera Baetidae) in KorearsquoThe Korean Journal of Systematics Biology 13 303ndash314

Bae YJ and Park SY (1998) lsquoAlainites Baetis Labiobaetis and Nigrobaetis(Ephemeroptera Baetidae) in Korearsquo The Korean Journal of Systematics Biology 141ndash12

Banks N (1900) lsquoNew genera and species of Nearctic Neuropteroid Insectsrsquo Transactions ofthe American Entomological Society 26 239ndash259

Banks N (1914) lsquoNew neuropteroid insects native and exoticrsquo Proceedings of the Academy ofNatural Sciences of Philadelphia 66 608ndash632

Aquatic Insects 53

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

After 150 years of intensive works the systematics of Baetidae can be consideredas relatively well known even if some taxomomic problems are still not completelysolved The validity of some species needs to be reviewed a few of them have neverbeen mentioned since their original description

Australasian realm

The Australasian realm presently encompasses 40 known species all of themendemic except for Cloeon virens which is also distributed in the Oriental realmTwelve genera occur in this area only six of them being endemic (50) (Table 1)The non-endemic genera are also distributed in the Oriental region The presence ofsome typical Palearctic genera such as Centroptilum in Australasia is doubtful andshould be clarified in the near future

Offadens soror was the first species described from this region by Ulmer in 1908Following Ulmer Tillyard and Harker provided the only significant works onAustralian Baetidae (Tillyard 1936 Harker 1950 1954 1957) and until recently(Figure 6) Suter greatly improved the knowledge of the Ephemeroptera fauna in theAustralian region providing the first key for the larval stage of mayflies (Suter 1979)At that time the Baetidae encompassed only 12 species and five genera andinterestingly Bungona was the only genus which was endemic from this area all theother Australian species being attributed to Palearctic or Holarctic genera Later onSuter described new species and redescribed some genera (Suter 1986 2000 2001Suter and Pearson 2001) He also provided a key specifically for the larvae ofBaetidae which included several undescribed species and genera (Suter 1997)

As in other biogeographic regions Lugo-Ortiz and McCafferty played animportant role by describing several species and four endemic genera from thisregion (Lugo-Ortiz and McCafferty 1998e 1999a 1999c) and by reporting generasuch as Cloeodes or LabiobaetisPseudocloeon (Lugo-Ortiz and McCafferty 1998bLugo-Ortiz et al 1999) They also described several new taxa from New Guinea(Lugo-Ortiz and McCafferty 1999c Lugo-Ortiz et al 1999) Baetidae were one of thetwo families of mayflies reported from the Fiji Islands although it was only apreliminary study the diversity was relatively high with 12 species and three genera(Flowers 1990)

Figure 6 Specific and generic diversity in Australasian realm Square total number ofspecies (left scale) Triangle total number of genera (right scale)

Aquatic Insects 51

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Although the predaceous habit is very unusual in Baetidae two of the 12 generaof this region are carnivorous Echinobaetis from Sulawesi (Mol 1989) andMystaxyops from New Guinea (McCafferty and Sun 2005)

New Zealand is the only important landmass where mayflies are present andBaetidae have never been found This gap remains an enigma These islands are quiteisolated and possess endemic families such as Siphlaenigmatidae and Rallidentidaebut also the widespread family Leptophlebiidae The Baetidae were also consideredto be absent from New Caledonia (Peters 2001) However in material recentlycollected in standing waters half a dozen larvae of Cloeon were included(unpublished data) Because of the absence of Baetidae in previous intensivesamplings and the high power of dispersion of Cloeon it seems likely that a recentcolonisation of the island may be related to human activity If the Leptophlebiidaewere able to remain established on these islands after the break-off of the Gondwanaor to colonise the islands afterwards why were the Baetidae not able to do the sameThe question remains open

Despite recent improvements the fauna of this region remains insufficientlyknown in Australia and even more so on other islands and archipelagos ForAustralia a recent estimation established that only about half of the Australianspecies of Baetidae are described and probably some new genera still need to beestablished (Jeff Webb personal communication)

The biogeographic affinities of this region need to be analysed in detail All thenon-endemic genera are also distributed in the Oriental region they are mainlydiversified in the North East of Australasia and had probably dispersed step by stepthrough South East Asia into Australia where their distribution is limited to theNorth (Lugo-Ortiz and McCafferty 1999b) The genus Cloeodes clearly connects thisarea with the history of Gondwana and the continents placed in the SouthHemisphere Endemic genera such Edmundsiops Offadens and Bungona may berelated to South American taxa (Jeff Webb personal communication)

Conclusion

Since the establishment of the Baetidae in 1815 their knowledge has been hugelyincreased due to the efforts of many authors around the world In 1999 Lugo-Ortizand McCafferty provided the first worldwide synthesis of this family They estimatedthe diversity of this family at 650 species and 87 genera (Lugo-Ortiz and McCafferty1999b) Ten years later the number of species has increased by almost one third toreach 894 species During these 10 years 142 new species were described mainlyfrom South America Madagascar and West Africa It means that the increase of thetotal number of species is mainly due to the description of new species but Lugo-Ortiz and McCafferty had also underestimated the number of species of Baetidae byabout 100 species During these 10 years 17 genera were described and four weresynonymised

Baetidae are mostly diversified and possess a much higher rate of genericendemism in the Tropics and Southern Hemisphere than in the NorthernHemisphere (Barber-James et al 2008) The Afrotropics and Neotropics possessby far the most diversified faunas Despite common Gondwanian origins thesefaunas have almost no affinities The Palearctic fauna shares most genera with theOriental andor Afrotropical faunas as well as the Nearctic fauna Stepwisecolonisation could have occurred through the Palearctic region Central America

52 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

constitutes a bridge between North and South Americas and is under the influence ofboth areas (McCafferty 1998) The affinities of the Nearctic and Neotropical faunasare mainly due to the presence of North American taxa in Central America TheOriental fauna is still only partially known regions of biogeographic importance andwith a high potential diversity such as the Indian subcontinent and South East Asiaremain almost completely unknown Local studies clearly indicate that the diversityis quite high but the distribution of the taxa remains unknown The Oriental realm istherefore probably the most promising and challenging area Even if the globaldiversity is probably lower in Australasia the lack of knowledge of the fauna isobvious especially in the tropical part of this area such as New Guinea

The knowledge on systematics and faunistic composition are the first inevitablesteps to understand and preserve a natural system Most of the remaining unknownor poorly known faunas are under a direct human threat This underlines theurgency of this work before these faunas are definitively lost this is especially true inmost of the tropical rain forest areas where a primordial aquatic fauna is still presentFor areas with sufficient knowledge it is also the duty of the systematists to offer thenecessary tools for a correct identification of the taxa We can only encourageeveryone to make our knowledge accessible to others scientists especially by creatingkeys and checklists at different systematic levels

A comprehensive reconstruction of the phylogeny of the Baetidae is also highlynecessary Such a reconstruction should be based on modern tools and include bothmolecular and morphological characters It is the only way to solve contradictorysuprageneric relationships Such studies would allow a more accurate understandingof the relationships between the different areas in a global biogeographic approachThey would help to understand the historical influence of Gondwana and Laurasiaon the mayfly fauna

Acknowledgements

We want to express our appreciation to Michel Sartori (Museum of Zoology LausanneSwitzerland) and Helen Barber-James (Albany Museum Grahamstown South Africa) foruseful comments on this paper and great help in the conception of the Baetidae databaseAline Pasche (Museum of Zoology Lausanne Switzerland) offered technical supportFinancial support from the Argentine National Council of Scientific and TechnologicalResearch (CONICET) and ANPCYT (Pict 524-Pict 528) to which the junior author belongsare acknowledged The electronic database lsquolsquoEphemeroptera of the Worldrsquorsquo of Nikita Kluge(httpwwwinsectabiopuruzEph-sppContentshtm) was of great help for cross-checkingof our own database

References

Agnew JD (1961) lsquoNew Baetidae (Ephem) from South Africarsquo Novos Taxa Entomologicos25 3ndash18

Agnew JD (1963) lsquoNew South African records of imperfectly known Baetidae (Ephem)rsquoHydrobiologia 22 41ndash46

Bae YJ (1997) lsquoTaxonomy of Cloeon and Procloeon (Ephemeroptera Baetidae) in KorearsquoThe Korean Journal of Systematics Biology 13 303ndash314

Bae YJ and Park SY (1998) lsquoAlainites Baetis Labiobaetis and Nigrobaetis(Ephemeroptera Baetidae) in Korearsquo The Korean Journal of Systematics Biology 141ndash12

Banks N (1900) lsquoNew genera and species of Nearctic Neuropteroid Insectsrsquo Transactions ofthe American Entomological Society 26 239ndash259

Banks N (1914) lsquoNew neuropteroid insects native and exoticrsquo Proceedings of the Academy ofNatural Sciences of Philadelphia 66 608ndash632

Aquatic Insects 53

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Although the predaceous habit is very unusual in Baetidae two of the 12 generaof this region are carnivorous Echinobaetis from Sulawesi (Mol 1989) andMystaxyops from New Guinea (McCafferty and Sun 2005)

New Zealand is the only important landmass where mayflies are present andBaetidae have never been found This gap remains an enigma These islands are quiteisolated and possess endemic families such as Siphlaenigmatidae and Rallidentidaebut also the widespread family Leptophlebiidae The Baetidae were also consideredto be absent from New Caledonia (Peters 2001) However in material recentlycollected in standing waters half a dozen larvae of Cloeon were included(unpublished data) Because of the absence of Baetidae in previous intensivesamplings and the high power of dispersion of Cloeon it seems likely that a recentcolonisation of the island may be related to human activity If the Leptophlebiidaewere able to remain established on these islands after the break-off of the Gondwanaor to colonise the islands afterwards why were the Baetidae not able to do the sameThe question remains open

Despite recent improvements the fauna of this region remains insufficientlyknown in Australia and even more so on other islands and archipelagos ForAustralia a recent estimation established that only about half of the Australianspecies of Baetidae are described and probably some new genera still need to beestablished (Jeff Webb personal communication)

The biogeographic affinities of this region need to be analysed in detail All thenon-endemic genera are also distributed in the Oriental region they are mainlydiversified in the North East of Australasia and had probably dispersed step by stepthrough South East Asia into Australia where their distribution is limited to theNorth (Lugo-Ortiz and McCafferty 1999b) The genus Cloeodes clearly connects thisarea with the history of Gondwana and the continents placed in the SouthHemisphere Endemic genera such Edmundsiops Offadens and Bungona may berelated to South American taxa (Jeff Webb personal communication)

Conclusion

Since the establishment of the Baetidae in 1815 their knowledge has been hugelyincreased due to the efforts of many authors around the world In 1999 Lugo-Ortizand McCafferty provided the first worldwide synthesis of this family They estimatedthe diversity of this family at 650 species and 87 genera (Lugo-Ortiz and McCafferty1999b) Ten years later the number of species has increased by almost one third toreach 894 species During these 10 years 142 new species were described mainlyfrom South America Madagascar and West Africa It means that the increase of thetotal number of species is mainly due to the description of new species but Lugo-Ortiz and McCafferty had also underestimated the number of species of Baetidae byabout 100 species During these 10 years 17 genera were described and four weresynonymised

Baetidae are mostly diversified and possess a much higher rate of genericendemism in the Tropics and Southern Hemisphere than in the NorthernHemisphere (Barber-James et al 2008) The Afrotropics and Neotropics possessby far the most diversified faunas Despite common Gondwanian origins thesefaunas have almost no affinities The Palearctic fauna shares most genera with theOriental andor Afrotropical faunas as well as the Nearctic fauna Stepwisecolonisation could have occurred through the Palearctic region Central America

52 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

constitutes a bridge between North and South Americas and is under the influence ofboth areas (McCafferty 1998) The affinities of the Nearctic and Neotropical faunasare mainly due to the presence of North American taxa in Central America TheOriental fauna is still only partially known regions of biogeographic importance andwith a high potential diversity such as the Indian subcontinent and South East Asiaremain almost completely unknown Local studies clearly indicate that the diversityis quite high but the distribution of the taxa remains unknown The Oriental realm istherefore probably the most promising and challenging area Even if the globaldiversity is probably lower in Australasia the lack of knowledge of the fauna isobvious especially in the tropical part of this area such as New Guinea

The knowledge on systematics and faunistic composition are the first inevitablesteps to understand and preserve a natural system Most of the remaining unknownor poorly known faunas are under a direct human threat This underlines theurgency of this work before these faunas are definitively lost this is especially true inmost of the tropical rain forest areas where a primordial aquatic fauna is still presentFor areas with sufficient knowledge it is also the duty of the systematists to offer thenecessary tools for a correct identification of the taxa We can only encourageeveryone to make our knowledge accessible to others scientists especially by creatingkeys and checklists at different systematic levels

A comprehensive reconstruction of the phylogeny of the Baetidae is also highlynecessary Such a reconstruction should be based on modern tools and include bothmolecular and morphological characters It is the only way to solve contradictorysuprageneric relationships Such studies would allow a more accurate understandingof the relationships between the different areas in a global biogeographic approachThey would help to understand the historical influence of Gondwana and Laurasiaon the mayfly fauna

Acknowledgements

We want to express our appreciation to Michel Sartori (Museum of Zoology LausanneSwitzerland) and Helen Barber-James (Albany Museum Grahamstown South Africa) foruseful comments on this paper and great help in the conception of the Baetidae databaseAline Pasche (Museum of Zoology Lausanne Switzerland) offered technical supportFinancial support from the Argentine National Council of Scientific and TechnologicalResearch (CONICET) and ANPCYT (Pict 524-Pict 528) to which the junior author belongsare acknowledged The electronic database lsquolsquoEphemeroptera of the Worldrsquorsquo of Nikita Kluge(httpwwwinsectabiopuruzEph-sppContentshtm) was of great help for cross-checkingof our own database

References

Agnew JD (1961) lsquoNew Baetidae (Ephem) from South Africarsquo Novos Taxa Entomologicos25 3ndash18

Agnew JD (1963) lsquoNew South African records of imperfectly known Baetidae (Ephem)rsquoHydrobiologia 22 41ndash46

Bae YJ (1997) lsquoTaxonomy of Cloeon and Procloeon (Ephemeroptera Baetidae) in KorearsquoThe Korean Journal of Systematics Biology 13 303ndash314

Bae YJ and Park SY (1998) lsquoAlainites Baetis Labiobaetis and Nigrobaetis(Ephemeroptera Baetidae) in Korearsquo The Korean Journal of Systematics Biology 141ndash12

Banks N (1900) lsquoNew genera and species of Nearctic Neuropteroid Insectsrsquo Transactions ofthe American Entomological Society 26 239ndash259

Banks N (1914) lsquoNew neuropteroid insects native and exoticrsquo Proceedings of the Academy ofNatural Sciences of Philadelphia 66 608ndash632

Aquatic Insects 53

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

constitutes a bridge between North and South Americas and is under the influence ofboth areas (McCafferty 1998) The affinities of the Nearctic and Neotropical faunasare mainly due to the presence of North American taxa in Central America TheOriental fauna is still only partially known regions of biogeographic importance andwith a high potential diversity such as the Indian subcontinent and South East Asiaremain almost completely unknown Local studies clearly indicate that the diversityis quite high but the distribution of the taxa remains unknown The Oriental realm istherefore probably the most promising and challenging area Even if the globaldiversity is probably lower in Australasia the lack of knowledge of the fauna isobvious especially in the tropical part of this area such as New Guinea

The knowledge on systematics and faunistic composition are the first inevitablesteps to understand and preserve a natural system Most of the remaining unknownor poorly known faunas are under a direct human threat This underlines theurgency of this work before these faunas are definitively lost this is especially true inmost of the tropical rain forest areas where a primordial aquatic fauna is still presentFor areas with sufficient knowledge it is also the duty of the systematists to offer thenecessary tools for a correct identification of the taxa We can only encourageeveryone to make our knowledge accessible to others scientists especially by creatingkeys and checklists at different systematic levels

A comprehensive reconstruction of the phylogeny of the Baetidae is also highlynecessary Such a reconstruction should be based on modern tools and include bothmolecular and morphological characters It is the only way to solve contradictorysuprageneric relationships Such studies would allow a more accurate understandingof the relationships between the different areas in a global biogeographic approachThey would help to understand the historical influence of Gondwana and Laurasiaon the mayfly fauna

Acknowledgements

We want to express our appreciation to Michel Sartori (Museum of Zoology LausanneSwitzerland) and Helen Barber-James (Albany Museum Grahamstown South Africa) foruseful comments on this paper and great help in the conception of the Baetidae databaseAline Pasche (Museum of Zoology Lausanne Switzerland) offered technical supportFinancial support from the Argentine National Council of Scientific and TechnologicalResearch (CONICET) and ANPCYT (Pict 524-Pict 528) to which the junior author belongsare acknowledged The electronic database lsquolsquoEphemeroptera of the Worldrsquorsquo of Nikita Kluge(httpwwwinsectabiopuruzEph-sppContentshtm) was of great help for cross-checkingof our own database

References

Agnew JD (1961) lsquoNew Baetidae (Ephem) from South Africarsquo Novos Taxa Entomologicos25 3ndash18

Agnew JD (1963) lsquoNew South African records of imperfectly known Baetidae (Ephem)rsquoHydrobiologia 22 41ndash46

Bae YJ (1997) lsquoTaxonomy of Cloeon and Procloeon (Ephemeroptera Baetidae) in KorearsquoThe Korean Journal of Systematics Biology 13 303ndash314

Bae YJ and Park SY (1998) lsquoAlainites Baetis Labiobaetis and Nigrobaetis(Ephemeroptera Baetidae) in Korearsquo The Korean Journal of Systematics Biology 141ndash12

Banks N (1900) lsquoNew genera and species of Nearctic Neuropteroid Insectsrsquo Transactions ofthe American Entomological Society 26 239ndash259

Banks N (1914) lsquoNew neuropteroid insects native and exoticrsquo Proceedings of the Academy ofNatural Sciences of Philadelphia 66 608ndash632

Aquatic Insects 53

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Banks N (1918) lsquoNew neuropteroid insectsrsquo Bulletin of the Museum of Comparative Zoology62(1) 1ndash22

Barber-James HM (2007) lsquoList of Afrotropical mayfly families genera and species withsynonymsrsquo httpoldwwwruaczaacademicdepartmentszooentoMartinEphemeropteraAfrica2007htm

Barber-James HM Gattolliat J-L Sartori M and Hubbard MD (2008) lsquoGlobaldiversity of mayflies (Ephemeroptera Insecta) in freshwaterrsquo Hydrobiologia 595339ndash350

Barber-James HM and Lugo-Ortiz CR (2003) lsquoEphemeropterarsquo in Guides to theFreshwater Invertebrates of Southern Africa Volume 7 Insecta 1 eds IJ de Moor JADay and FC de Moor Water Resource Commission Pretoria South Africa pp 16ndash159

Barnard KH (1932) lsquoSouth African may-flies (Ephemeroptera)rsquo Transactions of the RoyalSociety of South Africa 20 201ndash259

Bengtsson S (1912) lsquoNeue Ephemeriden aus Schwedenrsquo Entomologisk Tidskrift 33 107ndash117Bengtsson S (1914) lsquoBemerkungen uber die nordischen Arten der Gattung Cloeon Leachrsquo

Entomologisk Tidskrift 35 210ndash220Bengtsson S (1917) lsquoWeitere Beitrage zur Kenntnis der nordischen Eintagsfliegenrsquo

Entomologisk Tidskrift 38 174ndash194Bogoescu CD and Tabacaru I (1957) lsquoContributii la studiul sistematic al

nimfelor de Ephemeroptere din RPRrsquo Bulletin de la Section Scientifique de Roumanie9 241ndash284

Crass RS (1947) lsquoThe May-flies (Ephemeroptera) of Natal and the Eastern Capersquo Annals ofthe Natal Museum 11 37ndash110

Demoulin G (1964a) lsquoEphemeropterarsquo Parc National de lrsquoUpemba - Mission G F de Witte68 13ndash27

Demoulin G (1964b) lsquoMission H Loffler en Afrique orientale Ephemeropterarsquo Bulletin etAnnales de la Societe Entomologique de Belgique 100 279ndash294

Demoulin G (1965) lsquoEphemeropterarsquo Annales du Musee royal drsquoAfrique centrale 13891ndash114

Demoulin G (1966) lsquoQuelques Ephemeropteres nouveaux de Madagascarrsquo Annales de laSociete Entomologique de France 2 711ndash717

Demoulin G (1967) lsquoDescription de deux larves atypiques de Baetidae (InsEphemeroptera)rsquo Bulletin et Annales de la Societe Entomologique de Belgique 103226ndash233

Demoulin G (1968) lsquoQuelques Ephemeropteres nouveaux de Madagascar IIrsquo Bulletin delrsquoInstitut Royal des Sciences Naturelles de Belgique 44 1ndash9

Demoulin G (1970) lsquoEphemeroptera des faunes ethiopienne et malgachersquo South AfricanAnimal Life 14 24ndash170

Domınguez E Molineri C Pescador ML Hubbard MD and Nieto C (2006)Ephemeroptera of South America Sofia and Moscow Pensoft Press

Dominique Y Thomas A and Fenoglio S (2002) lsquoRedescription de Camelobaetidiusmusseri (Traver amp Edmunds 1968) (Ephemeroptera Baetidae)rsquo Ephemera 3 33ndash41

Dubey OP (1971) lsquoTorrenticole insects of the Himalaya VI Description of nine species ofEphemerida from the Northwest Himalayarsquo Oriental Insects 5 521ndash548

Eaton AE (1869) lsquoOn Centroptilum a new genus of the Ephemeridaersquo EntomologistrsquosMonthly Magazine 6 131ndash132

Eaton AE (1870) lsquoOn some new British species of Ephemeridaersquo Transactions of theEntomological Society of London 1870 1ndash8

Eaton AE (1871) lsquoA Monograph of the Ephemeridaersquo Transactions of the EntomologicalSociety of London 1871 1ndash164

Eaton AE (1881) lsquoAn announcement of new genera of the Ephemeridaersquo EntomologistMonthly Magazine 17 191ndash197

Eaton AE (1883ndash1888) lsquoA revisional monograph of recent Ephemeridae or mayfliesrsquoTransactions of the Linnean Society of London 2nd Ser Zoology 2 1ndash352

Edmunds GF (1954) lsquoThe Mayflies of Utahrsquo Proceedings of the Utah Academy of SciencesArts and Letters 31 64ndash66

Edmunds GF (1972) lsquoBiogeography and evolution of Ephemeropterarsquo Annual Review ofEntomology 17 21ndash42

54 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Edmunds GF (1975) lsquoPhylogenetic biogeography of mayfliesrsquo Annals of the MissouriBotanical Garden 62 251ndash263

Edmunds GF and Allen RK (1957a) lsquoA checklist of the Ephemeroptera ofNorth America North of Mexicorsquo Annals Of The Entomological Society of America 50317ndash324

Edmunds GF and Allen RK (1957b) lsquoA new species of Baetis from Oregon(Ephemeroptera Baetidae)rsquo Journal of the Kansas Entomological Society 30 57ndash58

Edmunds GF Jensen SL and Berner L (1976) The Mayflies of North and CentralAmerica Minneapolis MNUniversity of Minnesota Press

Elouard J-M and Gibon F-M (2001) Biodiversite et biotypologie des eaux continentalesmalgaches Montpellier IRD

Elouard J-M Gillies MT and Wuillot J (1990) lsquoEphemeroptera from West Africa Thegenus Pseudopannota (Baetidae)rsquo Revue drsquoHydrobiologie tropicale 23 27ndash39

Elouard J-M and Hideux P (1991) lsquoMayflies of West Africa Thraulobaetodes an atypicalnew genus of crawling Baetidaersquo in Overviews and strategies of the Ephemeroptera andPlecoptera eds J Alba-Tercedor and A Sanchez-Ortega Gainesville FLSandhill CranePress Inc pp 169ndash174

Esben-Petersen P (1913) lsquoEphemeridae from South Africarsquo Annals of the South AfricanMuseum 10 177ndash187

Flowers RW (1990) lsquoEphemeroptera on the Fiji Islandsrsquo in Mayflies and Stoneflies Lifestory and Biology ed IC Campbell Dordrecht Kluwer Academic Publisherspp 125ndash134

Fujitani T (2008) lsquoThe family Baetidae from Japanrsquo in International Advances in the EcologyZoogeography and Systematics of Mayflies and Stoneflies eds FR Hauer JA Stanfordand RL Newell Berkeley University of California Press pp 205ndash218

Fujitani T Hirowatari T and Tanida K (2003a) lsquoGenera and species of Baetidae in JapanNigrobaetis Alainites Labiobaetis and Tenuibaetis n stat (Ephemeroptera)rsquo Limnology4 121ndash129

Fujitani T Hirowatari T and Tanida K (2003b) lsquoNymphs of Nigrobaetis AlainitesLabiobaetis Tenuibaetis and Baetis from Japan (Ephemeroptera Baetidae) Diagnosesand keys for genera and speciesrsquo in Research Update on Ephemeroptera and Plecoptera edE Gaino Perugia University of Perugia Italy pp 127ndash133

Gattolliat J-L (2001a) lsquoThe genus Cloeodes (Ephemeroptera Baetidae) in MadagascarrsquoRevue Suisse de Zoologie 108 387ndash402

Gattolliat J-L (2001b) lsquoRheoptilum a new genus of two-tailed Baetidae (Ephemeroptera)from Madagascarrsquo Aquatic Insects 23 67ndash81

Gattolliat J-L (2002a) lsquoThree new Malagasy species of Xyrodromeus (EphemeropteraBaetidae) with the first generic description of the adultsrsquo Revue Suisse de Zoologie 109325ndash341

Gattolliat J-L (2002b) lsquoTwo new genera of Baetidae (Ephemeroptera Insecta) fromMadagascarrsquo Aquatic Insects 24 143ndash159

Gattolliat J-L (2004) lsquoA distinctive new species of Xyrodromeus Lugo-Ortiz amp McCafferty(Ephemeroptera Baetidae) from Madagascarrsquo Zootaxa 452 1ndash10

Gattolliat JL Monaghan MT Sartori M Elouard JM James H Derleth P GlaizotO de Moor F and Vogler AP (2008) lsquoA molecular analysis of the AfrotropicalBaetidaersquo in International Advances in the Ecology Zoogeography and Systematics ofMayflies and Stoneflies eds FR Hauer JA Stanford and RL Newell BerkeleyUniversity of California Press pp 219ndash232

Gattolliat J-L and Rabeantoandro SZ (2002) lsquoThe genus Cloeon (EphemeropteraBaetidae) in Madagascarrsquo Mitteilungen der Schweizerischen entomologischen Gesellschaft74 195ndash209 (2001)

Gattolliat J-L and Sartori M (2000) lsquoContribution to the systematics of the genusDabulamanzia (Ephemeroptera Baetidae) in Madagascarrsquo Revue Suisse de Zoologie 107561ndash577

Gattolliat J-L and Sartori M (2001) lsquoPredaceous Baetidae in Madagascar anuncommon and unsuspected high diversityrsquo in Trends in Research in Ephemeroptera andPlecoptera ed E Domınguez New York Kluwer AcademicPlenum Publisherspp 321ndash330

Aquatic Insects 55

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Gattolliat J-L and Sartori M (2003) lsquoAn overview of the Baetidae of Madagascarrsquo inResearch Update on Ephemeroptera and Plecoptera ed E Gaino Perugia University ofPerugia Italy pp 135ndash144

Gattolliat J-L and Sartori M (2008) lsquoOrder Ephemeropterarsquo in Arthropod Fauna of theUAE ed A van Harten Abu Dhabi Dar Al Ummah pp 47ndash83

Gillies MT (1949) lsquoNotes on some Ephemeroptera Baetidae from India and South-EastAsiarsquo Transactions of the Royal Entomological Society of London 100 161ndash177

Gillies MT (1985) lsquoA preliminary account of the East African species of Cloeon Leach andRhithrocloeon gen n (Ephemeroptera)rsquo Aquatic Insects 7 1ndash17

Gillies MT (1988) lsquoDescription of the nymphs of some Afrotropical Baetidae (Ephemer-optera) I Cloeon Leach and Rhithrocloeon Gilliesrsquo Aquatic Insects 10 49ndash59

Gillies MT (1990) lsquoA revision of the African species of Centroptilum Eaton (BaetidaeEphemeroptera)rsquo Aquatic Insects 12 97ndash128

Gillies MT (1991a) lsquoA diphyletic origin for the two-tailed baetid nymphs occurring inEast African stony streams with a description of the new genus and species Tanzaniellaspinosa gen sp novrsquo in Overview and strategies of Ephemeroptera and Plecoptera edsJ Alba-Tercedor and A Sanchez-Ortega Gainesville FL Sandhill Crane Press Incpp 175ndash187

Gillies MT (1991b) lsquoA new species of Afroptilum (Afroptiloides) from East Africa (EphemBaetidae)rsquo Entomologistrsquos Monthly Magazine 127 109ndash115

Gillies MT (1994) lsquoDescriptions of some Afrotropical Baetidae (Ephemeroptera) II BaetisLeach sl East African speciesrsquo Aquatic Insects 16 105ndash118

Gillies MT (1998) lsquoThe mayflies (Ephemeroptera) of the upper River Sigi North-EastTanzaniarsquo Freshwater Forum 10 49ndash57

Gillies MT and Elouard J-M (1990) lsquoThe mayfly-mussel association a new example fromthe River Niger Basinrsquo in Mayflies and Stoneflies Life story and Biology ed ICCampbell Dordrecht Kluwer Academic Publishers pp 289ndash298

Gillies MT Elouard J-M and Wuillot J (1990) lsquoEphemeroptera from West Africa thegenus Ophelmatostoma (Baetidae)rsquo Revue drsquoHydrobiologie Tropicale 23(2) 115ndash120

Gillies MT and Wuillot J (1997) lsquoPlatycloeon a new genus of Baetidae (Ephemeroptera)from East Africarsquo Aquatic Insects 19 185ndash189

Glazaczow A (1997) lsquoObservations on the Psammophilous mayfly species Procleon nanumin the North East of Polandrsquo in Ephemeroptera amp Plecoptera Biology-Ecology-Systematics eds P Landolt and M Sartori Fribourg Mauron thorn Tinguely amp LachatSA pp 83ndash87

Gose K (1980) lsquoNihon san kagero-rui [The mayflies of Japanese Key to families genera andspecies]rsquo (in Japanese) Aquabiology (Nara) 2(1) 76ndash79 2(2) 122ndash123 2(3) 211ndash2152(4) 286ndash288 2(5) 366ndash368 2(6) 454ndash457

Hagen H (1858) lsquoSynopsis der Neuroptera Ceylonsrsquo Verhandlungen der Kaiserlich-Koniglichen Zoologisch-Botanischen Gesellschaft in Wien 8 471ndash488

Hagen H (1861) lsquoSynopsis of the Neuroptera of North America with a list of the SouthAmerican speciesrsquo Smithsonian Miscellaneous collections 1ndash347

Harker JE (1950) lsquoAustralian Ephemeroptera Part I Taxonomy of New South Walesspecies and evaluation of taxonomic charactersrsquo Proceedings of the Linnean Society ofNew South Wales 75 1ndash34

Harker JE (1954) lsquoThe Ephemeroptera from Eastern Australiarsquo Transactions of the RoyalEntomological Society of London 105 241ndash268

Harker JE (1957) lsquoSome new Australian Ephemeropterarsquo Proceedings of the RoyalEntomological Society of London B 26 63ndash78

Jacobus LM and McCafferty WP (2005a) lsquoApobaetis futilis (McDunnough) a newcombination in Nearctic Baetidae (Ephemeroptera)rsquo Proceedings of the EntomologicalSociety of Washington 107 979

Jacobus LM and McCafferty WP (2005b) lsquoA new species and new synonymin Camelobaetidius Demoulin (Ephemeroptera Baetidae)rsquo Journal of the KansasEntomological Society 78 153ndash157

Jacobus LM and McCafferty WP (2006) lsquoA new species of Acentrella Bengtsson(Ephemeroptera Baetidae) from Great Smoky Mountains National Park USArsquo AquaticInsects 28 101ndash111

56 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Kang SC Chang HC and Yang CT (1994) lsquoA revision of the genus Baetis in Taiwan(Ephemeroptera Baetidae)rsquo Journal of Taiwan Museum 47 9ndash44

Kang SC and Yang CT (1996) lsquoTwo new species of Baetis Leach (EphemeropteraBaetidae) from Taiwanrsquo Chinese Journal of Entomology 16 61ndash66

Kapur AP and Kripalani MB (1963) lsquoThe mayflies (Ephemeroptera) from the north-western Himalayarsquo Records of the Indian Museum 59 183ndash221 (1961)

Kazlauskas R (1963) lsquoNew and little known may flies (Ephemeroptera) from the USSRrsquoRevue drsquoEntomologie de lrsquoURSS 42 582ndash593

Kimmins DE (1947) lsquoNew species of Indian Ephemeropterarsquo Proceedings of the RoyalEntomological Society of London B 16 92ndash100

Kimmins DE (1956) lsquoNew species of Ephemeroptera from Ugandarsquo Bulletin of the BritishMuseum (Natural History) Entomology 4 71ndash87

Kopelke J-P (1980) lsquoEphemeroptera aus der Emergenz des zentralafrikanischen BergbachesKalengo (Zaıre) I Baetidaersquo Entomologische Abhandlungen 43 99ndash129

Kluge N Ju (2009) lsquoNew version of the database lsquolsquoEphemeroptera of the Worldrsquorsquo as the firstexperience of a permanent and objective web catalogue in biologyrsquo in InternationalPerspectives in Mayfly and Stonefly Research Proceedings of the 12th InternationalConference on Ephemeroptera and the 16th International Symposium on PlecopteraStuttgart 2008 ed AH Staniczek Aquatic Insects 31 Supplement 1 167ndash180

Lestage J-A (1918) lsquoLes Ephemeres drsquoAfrique (notes critiques sur les especes connues)rsquoRevue zoologique africaine 6 65ndash114

Leveque C Hougard JM Resh V Statzner B and Yameogo L (2003) lsquoFreshwaterecology and biodiversity in the tropics what did we learn from 30 years of onchocerciasiscontrol and the associated biomonitoring of West African riversrsquo Hydrobiologia 50023ndash49

Lugo-Ortiz CR (1999) lsquoSystematic studies of Baetidae (Insecta Ephemeroptera) withemphasis on biodiversity in the Southern Hemispherersquo Purdue University Department ofEntomology

Lugo-Ortiz CR and McCafferty WP (1995) lsquoThree distinctive new genera of Baetidae(Insecta Ephemeroptera) from South Americarsquo Annales de Limnologie 31 233ndash243

Lugo-Ortiz CR and McCafferty WP (1996a) lsquoAturbina georgei gen et sp n A smallminnow mayfly (Ephemeroptera Baetidae) without turbinate eyesrsquo Aquatic Insects 18175ndash183

Lugo-Ortiz CR and McCafferty WP (1996b) lsquoThe Bugilliesia complex of AfricanBaetidae (Ephemeroptera)rsquo Transactions of the American Entomological Society 122175ndash197

Lugo-Ortiz CR and McCafferty WP (1997) lsquoNew Afrotropical genus of Baetidae(Insecta Ephemeroptera) with bladelike mandiblesrsquo Bulletin de la Societe drsquoHistoireNaturelle de Toulouse 133 41ndash46

Lugo-Ortiz CR and McCafferty WP (1998a) lsquoThe Centroptiloides Complex ofAfrotropical small minnow mayflies (Ephemeroptera Baetidae)rsquo Annals of TheEntomological Society of America 91 1ndash26

Lugo-Ortiz CR and McCafferty WP (1998b) lsquoFirst report and new species of the genusCloeodes (Ephemeroptera Baetidae) from Australiarsquo Entomological News 109 122ndash128

Lugo-Ortiz CR and McCafferty WP (1998c) lsquoFive new genera of Baetidae (InsectaEphemeroptera) from South Americarsquo Annales de Limnologie 34 57ndash73

Lugo-Ortiz CR and McCafferty WP (1998d) lsquoA new North American genus ofBaetidae (Ephemeroptera) and key to Baetis complex generarsquo Entomological News 109345ndash353

Lugo-Ortiz CR and McCafferty WP (1998e) lsquoOffadens a new genus of small minnowmayflies (Ephemeroptera Baetidae) from Australiarsquo Proceedings of the EntomologicalSociety of Washington 100 306ndash309

Lugo-Ortiz CR and McCafferty WP (1998f) lsquoPhylogeny and biogeography of Nesydemiusn gen and related Afrotropical genera (Insecta Ephemeroptera Baetidae)rsquo Bulletin de laSociete drsquoHistoire Naturelle de Toulouse 134 7ndash12

Lugo-Ortiz CR and McCafferty WP (1999a) lsquoEdmundsiops instigatus A new genusand species of small minnow mayflies (Ephemeroptera Baetidae) from AustraliarsquoEntomological News 110 65ndash69

Aquatic Insects 57

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Lugo-Ortiz CR and McCafferty WP (1999b) lsquoGlobal biodiversity of the mayfly familyBaetidae (Ephemeroptera) a generic perspectiversquo Trends in Entomology 2 45ndash54

Lugo-Ortiz CR and McCafferty WP (1999c) lsquoA new genus of small minnow mayflies(Ephemeroptera Baetidae) with six new species from New Guinea and New BritainrsquoAnnales de Limnologie 35 57ndash70

Lugo-Ortiz CR and McCafferty WP (1999d) lsquoRevision of South American species ofBaetidae (Ephemeroptera) previously placed in Baetis Leach and Pseudocloeon KlapalekrsquoAnnales de Limnologie 35 257ndash262

Lugo-Ortiz CR and McCafferty WP (1999e) lsquoThree new genera of small minnow mayflies(Ephemeroptera Baetidae) from the Andes and Patagoniarsquo Studies on Neotropical Faunaand Environment 34 88ndash104

Lugo-Ortiz CR McCafferty WP and Waltz RD (1999) lsquoDefinition and reorganizationof the genus Pseudocloeon (Ephemeroptera Baetidae) with new species descriptions andcombinationsrsquo Transactions of the American Entomological Society 125 1ndash37

Mayo VK (1973) lsquoFour news species of the genus Baetodes (Ephemeroptera Baetidae)rsquo ThePan-Pacific Entomologist 49 308ndash314

McCafferty WP (1998) lsquoEphemeroptera and the great American interchangersquo Journal of theNorth American Benthological Society 17 1ndash20

McCafferty WP (2000) lsquoNotations on South American Baetidae (Ephemeroptera)rsquoEntomological News 111 375ndash379

McCafferty WP (2009) lsquoMayfly Centralrsquo wwwentmpurdueeduEntomologyresearchmayflymayflyhtml

McCafferty WP and Baumgardner DE (2003) lsquoLugoiops maya a new genus and species ofEphemeroptera (Baetidae) from Central Americarsquo Proceedings of the EntomologicalSociety of Washington 105 397ndash406

McCafferty WP and Sun L (2005) lsquoMystaxiops A new genus of small minnow mayflies(Ephemeroptera Baetidae) from Papua New Guinearsquo Proceedings of the EntomologicalSociety of Washington 107 536ndash542

McDunnough J (1922) lsquoTwo new Canadian May-flies (Ephemeridae)rsquo Canadian Entomol-ogist 53 117ndash120

McDunnough J (1923) lsquoNew Canadian Ephemeridae with notesrsquo Canadian Entomologist55 39ndash50

McDunnough J (1924) lsquoNew Ephemeridae from Illinoisrsquo Canadian Entomologist 56 7ndash9McDunnough J (1925) lsquoNew Canadian Ephemeridae with notes IIIrsquo Canadian Entomol-

ogist 57(168ndash176) 185ndash192McDunnough J (1926) lsquoNew Canadian Ephemeridae with notes IVrsquo Canadian Entomol-

ogist 58 296ndash303McDunnough J (1929) lsquoNotes on North American Ephemeroptera with descriptions of new

species IIrsquo Canadian Entomologist 61 169ndash180McDunnough J (1931) lsquoNew species of North American Ephemeropterarsquo Canadian

Entomologist 63 82ndash93McDunnough J (1932) lsquoNew species of North American Ephemeroptera IIrsquo Canadian

Entomologist 64 209ndash215McDunnough J (1936) lsquoA new Arctic baetid (Ephemeroptera)rsquo Canadian Entomologist 68

32ndash34McDunnough J (1939) lsquoNew British Columbian Ephemeropterarsquo Canadian Entomologist

71 49ndash54Mol AWM (1986) lsquoHarpagobaetis gulosus gen nov sp nov a new mayfly from Suriname

(Ephemeroptera Baetidae)rsquo Zoologische Mededelingen 60 63ndash70Mol AWM (1989) lsquoEchinobaetis phagas gen nov spec nov a new mayfly from Sulawesi

(Ephemeroptera Baetidae)rsquo Zoologische Mededelingen 63 61ndash72Monaghan MT Gattolliat JL Sartori M Elouard JM James H Derleth P Glaizot

O de Moor F and Vogler AP (2005) lsquoTrans-oceanic and endemic origins of the smallminnow mayflies (Ephemeroptera Baetidae) of Madagascarrsquo Proceedings of The RoyalSociety B-Biological Sciences 272 1829ndash1836

Muller-Liebenau I (1980a) lsquoJubabaetis gen n and Platybaetis gen n two new genera of thefamily Baetidae from the Oriental Regionrsquo in Advances in Ephemeroptera Biology edsJF Flannagan and KE Marshall New York Plenum Press pp 103ndash114

58 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Muller-Liebenau I (1980b) lsquoA new species of the genus Platybaetis Muller-Liebenau 1980P bishopi spn from Malaysia (Insecta Ephemeroptera)rsquo Gewasser und Abwasser 666795ndash101

Muller-Liebenau I (1981) lsquoReview of the original material of the baetid genera Baetis andPseudocloeon from the Sunda Islands and the Philippines described by G Ulmer withsome general remarks (Insecta Ephemeroptera)rsquo Mitteilungen der hamburger zoologischerMuseum und Institut 78 197ndash208

Muller-Liebenau I (1982a) lsquoFive new species of Pseudocloeon Klapalek 1905 (FamBaetidae) from the Oriental region (Insecta Ephemeroptera) with some general remarkson Pseudocloeonrsquo Archiv fur Hydrobiologie 95 283ndash298

Muller-Liebenau I (1982b) lsquoA new genus and species of Baetidae from Sri Lanka (Ceylon)Indocloeon primum gen n sp n (Insecta Ephemeroptera)rsquo Aquatic Insects 4 125ndash129

Muller-Liebenau I (1982c) lsquoNew species of the family Baetidae from the Philippines (InsectaEphemeroptera)rsquo Archiv fur Hydrobiologie 94 70ndash82

Muller-Liebenau I (1983) lsquoThree new species of the genus Centroptella Braasch ampSoldan 1980 from Sri Lanka (Insecta Ephemeroptera)rsquo Archiv fur Hydrobiologie 97486ndash500

Muller-Liebenau I (1984a) lsquoBaetidae from Sabah (East Malaysia) (Ephemeroptera)rsquo inProceeding of the fourth international conference on Ephemeroptera eds V LandaT Soldan and M Tonner Bechyne CSAV pp 85ndash99

Muller-Liebenau I (1984b) lsquoNew genera and species of the family Baetidae from West-Malaysia (River Gombak) (Insecta Ephemeroptera)rsquo Spixiana 7 253ndash284

Muller-Liebenau I (1985) lsquoBaetidae from Taiwan with remarks on Baetiella Ueno 1931(Insecta Ephemeroptera)rsquo Archiv fur Hydrobiologie 104 93ndash110

Muller-Liebenau I and Heard WH (1979) lsquoSymbiocloeon a new genus of Baetidaefrom Thailand (Insecta Ephemeroptera)rsquo in Proceedings of the Second InternationalConference on Ephemeroptera eds K Pasternak and R Sowa Warszawa PanstwoweWydawnictwo Naukowe pp 57ndash66

Muller-Liebenau I and Hubbard MD (1985) lsquoBaetidae from Sri Lanka with some generalremarks on the Baetidae of the Oriental region (Insecta Ephemeroptera)rsquo The FloridaEntomologist 68 537ndash561

Muller-Liebenau I and Morihara DK (1982) lsquoIndobaetis A new genus of Baetidae fromSri Lanka (Insecta Ephemeroptera) with two new speciesrsquo Gewasser und Abwasser 686926ndash34

Navas L (1911) lsquoNotas entomologicas Excursiones por los alrededores de Granadarsquo Boletinde la Sociedad Aragonesa de Ciencias Naturales 10 204ndash211

Navas L (1913) lsquoNotas entomologicas Excursiones por los alrededores de Zaragozarsquo Boletinde la Sociedad Aragonesa de Ciencias Naturales 12 75ndash77

Navas L (1917) lsquoNeuropteros nuevos o poco concocidoslsquo Memorias de la Real Academia deCiencias y Artes de Barcelona 13 393ndash395

Navas L (1930) lsquoInsectes du Congo Belge (Serie IV)rsquo Revue de Zoologie et de BotaniqueAfricaines 19 305ndash336

Navas L (1931a) lsquoInsectes du Congo Belge (Serie V)rsquo Revue de Zoologie et de BotaniqueAfricaines 20 257ndash279

Navas L (1931b) lsquoNevropteres et insectes voisins - Chine et pays environnants - 2e SeriersquoNotes drsquoentomologie chinoise 1 1ndash10

Needham JG Traver JR and Hsu Y-S (1935) The biology of mayflies with a systematicaccount of North American species Ithaca NY Comstock Publishing Company

Nieto C (2003a) lsquoEl genero Camelobaetidius (Ephemeroptera Baetidae) en la ArgentinarsquoActa Zoologica Mexicana 88 233ndash255

Nieto C (2003b) lsquoA new species of Guajirolus (Ephemeroptera Baetidae) from Argentinaand description of a new genus from Boliviarsquo in Research Update on Ephemeroptera andPlecoptera ed E Gaino Perugia University of Perugia Italy pp 153ndash158

Nieto C (2004a) lsquoThe genus Baetodes (Ephemeroptera Baetidae) in South America with thedescription of new species from Argentina Bolivia and Perursquo Studies on NeotropicalFauna and Environment 39 63ndash79

Nieto C (2004b) lsquoRedescription of Varipes (Ephemeroptera Baetidae) with the descriptionof new species from Bolivia and Argentinarsquo Aquatic Insects 26 161ndash173

Aquatic Insects 59

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Nieto C (2004c) lsquoSouth American Baetidae (Ephemeroptera) a new generic synonymyrsquoStudies on Neotropical Fauna and Environment 39 95ndash101

Nieto C and Richard B (2008) lsquoThe genus Cloeodes (Ephemeroptera Baetidae) inArgentina with new generic synonymy and new speciesrsquo Zootaxa 1727 1ndash21

Nieto C and Salles FF (2006) lsquoRevision of the genus Paracloeodes (EphemeropteraBaetidae) in South Americarsquo Zootaxa 1ndash33

Nolte U Tietbohl RS and McCafferty WP (1996) lsquoA mayfly from tropical Brazilcapable of tolerating short-term dehydrationrsquo Journal of the North American BenthologicalSociety 15 87ndash94

Park SY Bae YJ and Yoon IB (1996) lsquoRevision of the Baetidae (Ephemeroptera) ofKorea (1) Historical review Acentrella Bengtsson and Baetiella Uenorsquo EntomologicalResearch Bulletin 22 55ndash66

Peters WL (2001) lsquoThe Ephemeroptera of New Caledonia and New Zealandrsquo in Trends inResearch in Ephemeroptera and Plecoptera ed E Domınguez New York KluwerAcademicPlenum Publishers pp 43ndash45

Pictet F-J (1843ndash1845) Histoire naturelle des Insectes Nevropteres Famille des EphemerinesGeneve

Rambur P (1842) Histoire Naturelle des Insectes Nevropteres Paris Librairie Encyclope-dique de Roret

Salles FF (2007) lsquoThe presence of Chane Nieto and Guajirolus Flowers (EphemeropteraBaetidae) in Brazil with the description of a new speciesrsquo Revista Brasileira DeEntomologia 51 404ndash409

Salles FF Andrade MB and Da-Silva ER (2005) lsquoCamelobaetidius francischettii a newspecies of Baetidae (Ephemeroptera) from Brazilrsquo Zootaxa 47ndash53

Salles FF and Batista JD (2004) lsquoThe presence of Varipes Lugo-Ortiz amp McCafferty(Ephemeroptera Baetidae) in Brazil with the description of a new speciesrsquo Zootaxa 4561ndash6

Salles FF Batista JD and Soares Cabette HR (2004) lsquoBaetidae (Insecta Ephemer-optera) de Nova Xavantina Mato Grosso Brasil novos registros e descricao de una novaespecie de Cloeodes Traverrsquo Biota Neotropica 4 1ndash8

Salles FF and Francischetti CN (2004) lsquoCryptonympha dasilvai sp nov (EphemeropteraBaetidae) do Brasilrsquo Neotropical Entomology 33 213ndash216

Salles FF and Polegatto CM (2008) lsquoTwo new species of Baetodes Needham amp Murphy(Ephemeroptera Baetidae) from Brazilrsquo Zootaxa 43ndash50

Sartori M Derleth P and Gattolliat JL (2003) lsquoNew data about the mayflies(Ephemeroptera) from Borneorsquo in Research Update on Ephemeroptera and Plecopteraed E Gaino Perugia University of Perugia Italy pp 403ndash406

Soldan T and Godunko RJ (2006) lsquoBaetis atlanticus n sp a new species of the subgenusRhodobaetis Jacob 2003 from Madeira Portugal (Ephemeroptera Baetidae)rsquo Genus 175ndash17

Soldan T and Thomas AGB (1983a) lsquoBaetis numidicus nsp Ephemeroptere nouveaudrsquoAlgerie (Baetidae)rsquo Annales de Limnologie 19 207ndash211

Soldan T and Thomas AGB (1983b) lsquoNew and little-known species of mayflies(Ephemeroptera) from Algeriarsquo Acta Entomologica Bohemoslovaca 80 356ndash376

Soldan T and Thomas AGB (1985) lsquoCentroptilum dimorphicum sp n a new species ofmayfly (Ephemeroptera Baetidae) from Algeriarsquo Acta Entomologica Bohemoslovaca 82180ndash186

Suter PJ (1979) lsquoA revised key to the Australian genera of mature mayfly(Ephemeroptera) nymphsrsquo Transactions of the Royal Society of South Australia 103 79ndash83

Suter PJ (1986) lsquoThe Ephemeroptera (Mayflies) of South Australiarsquo Records of the SouthAustralian Museum 19 339ndash397

Suter PJ (1997) lsquoPreliminary guide to the identification of nymphs of Australian Baetidmayflies (Insecta Ephemeroptera) found in flowing watersrsquo in Identification Guide No 14Albany Co-Operative Research Center for Freshwater Ecology

Suter PJ (2000) lsquoEdmundsiops hickmani spnov Offadens frater (Tillyard) nov comband description of the nymph of Cloeon tasmaniae Tillyard (EphemeropteraBaetidae) from Tasmaniarsquo Papers and Proceedings of the Royal Society of Tasmania134 63ndash73

60 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Suter PJ (2001) lsquoPlatybaetis gagadjuensis a new species from Northern Australia(Ephemeroptera Baetidae)rsquo in Trends in Research in Ephemeroptera and Plecopteraed E Dominguez New York Kluwer AcademicPlenum Publishers pp 359ndash364

Suter PJ and Pearson MJ (2001) lsquoRedescription of Bungona Harker with new synonymsin the Australian Baetidae (Insecta Ephemeroptera)rsquo Memoirs of the Museum of Victoria58 247ndash254

Thomas A and Gagneur J (1994) lsquoComplements et corrections a la faune desEphemeropteres drsquoAfrique du Nord 6 Alainites sadati n sp drsquoAlgerie (EphemeropteraBaetidae)rsquo Bulletin de la Societe drsquoHistoire Naturelle de Toulouse 130 43ndash45

Thomas A and Peru N (2003) lsquoLes Ephemeres de la Guyane Francaise 6 Description deWaltzoyphius roberti n sp (Ephemeroptera Baetidae)rsquo Bulletin de la Societe drsquoHistoireNaturelle de Toulouse 138 15ndash20 (2002)

Thomas AGB (1986) lsquoEphemeropteres du Sud-Ouest de la France IV Baetis catharus nspdes Pyrenees (Baetidae)rsquo Bulletin de la Societe drsquoHistoire Naturelle de Toulouse 122175ndash179

Thomas AGB (1998) lsquoA provisional checklist of the mayflies of North Africa(Ephemeroptera)rsquo Bulletin de la Societe drsquoHistoire Naturelle de Toulouse 134 13ndash20

Thomas AGB and Dia A (1984) lsquoBaetis baroukianus nsp Ephemeroptere nouveau duLiban (Baetidae)rsquo Bulletin de la Societe drsquoHistoire Naturelle de Toulouse 120 7ndash12

Thomas AGB and Dia A (1985) lsquoBaetis spei nsp Ephemeroptere nouveau du Libandescription et ecologie (Baetidae)rsquo Annales de Limnologie 21 241ndash245

Thomas AGB and Gazagnes G (1984) lsquoBaetis cyrneus n sp Ephemeroptere nouveau deCorse (Baetidae)rsquo Annales de Limnologie 20 199ndash202

Thomas AGB and Sartori M (1989) lsquoMayflies (Insecta Ephemeroptera) of SaudiArabiarsquo Fauna of Saudi Arabia 10 87ndash94

Thomas AGB and Soldan T (1987) lsquoBaetis ingridae n sp Ephemeroptere nouveau deCorse (Baetidae)rsquo Annales de Limnologie 23 23ndash26

Tillyard RJ (1936) lsquoThe trout-food insects of Tasmania Part II- A monograph of themayflies of Tasmaniarsquo Papers and proceedings of the Royal Society of Tasmania 193523ndash59

Tong X and Dudgeon D (2000) lsquoBaetiella (Ephemeroptera Baetidae) in Hong Kong withdescription of a new speciesrsquo Entomological News 111 143ndash148

Tong X and Dudgeon D (2003) lsquoFirst record of the genus Chopralla (EphemeropteraBaetidae) from China and description of a new speciesrsquo The Raffles Bulletin of Zoology51 17ndash19

Traver JR and Edmunds GF (1968) lsquoA revision of the Baetidae with spatulate-clawednymphs (Ephemeroptera)rsquo Pacific Insects 10 629ndash677

Ulmer G (1909) lsquoEphemeriden von Madagaskar und den Comorenrsquo Voeltzkowrsquos Reise inOstrafrika 1903ndash1905 II 365ndash368

Ulmer G (1939) lsquoEintagsfliegen (Ephemeropteren) von den Sunda-Inselnrsquo Archiv furHydrobiologie 16 443ndash692

Walsh BD (1863) lsquoObservations on certain N A Neuroptera by H Hagen M D ofKoenigsberg Prussia translated from the original French ms and published bypermission of the author with notes and descriptions of about twenty new N A speciesof Pseudoneuropterarsquo Proceedings of the Entomological Society of Philadelphia 2167ndash272

Walsh BD (1863) lsquoList of the Pseudoneuroptera of Illinois contained in the cabinet of thewriter with descriptions of over forty new species and notes on their structural affinitiesrsquoProceedings of the Academy of Natural Sciences of Philadelphia 1862 361ndash402

Waltz RD and McCafferty WP (1985) lsquoMoribaetis a new genus of NeotropicalBaetidae (Ephemeroptera)rsquo Proceedings of the Entomological Society of Washington 87239ndash251

Waltz RD and McCafferty WP (1986) lsquoApobaetis etowah (Traver) a new combination inNearctic Baetidae (Ephemeroptera)rsquo Proceedings of the Entomological Society ofWashington 88 191

Waltz RD and McCafferty WP (1987a) lsquoGeneric revision of Cloeodes and description oftwo new genera (Ephemeroptera Baetidae)rsquo Proceedings of the Entomological Society ofWashington 89 177ndash184

Aquatic Insects 61

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Waltz RD and McCafferty WP (1987b) lsquoNew genera of Baetidae for some Nearcticpreviously included in Baetis Leach (Ephemeroptera)rsquo Annals of the Entomological Societyof America 80 667ndash670

Waltz RD and McCafferty WP (1987c) lsquoSystematics of Pseudocloeon AcentrellaBaetiella and Liebebiella new genus (Ephemeroptera Baetidae)rsquo Journal of New YorkEntomological Society 95 553ndash568

Waltz RD and McCafferty WP (1989) lsquoNew species redescriptions and cladisticsof the genus Pseudocentroptiloides (Epherneroptera Baetidae)rsquo Journal of New YorkEntomological Society 97 151ndash158

Waltz RD McCafferty WP and Thomas A (1994) lsquoSystematics of Alainites n genDiphetor Indobaetis Nigrobaetis n stat and Takobia n stat (Ephemeroptera Baetidae)rsquoBulletin de la Societe drsquoHistoire Naturelle de Toulouse 130 33ndash36

Wang T-Q and McCafferty WP (1996) lsquoNew diagnostic characters for the mayfly familyBaetidae (Ephemeroptera)rsquo Entomological News 107 207ndash212

Weyenbergh H (1883) lsquoBijdrage tot de Kennis der zuid-amerikaansche EphemeridenrsquoTijdschrift voor Entomologie 26 159ndash174

Wiersema NA (2000) lsquoA new combination for two North American small minnow mayflies(Ephemeroptera Baetidae)rsquo Entomological News 111 140ndash142

Wiersema NA and McCafferty WP (2004) lsquoNew specific synonyms and records of NorthAmerican Centroptilum and Procloeon (Ephemeroptera Baetidae)rsquo Entomological News115 121ndash128

Wiersema NA Nelson CR and Kuehnl KF (2004) lsquoA new small minnow mayfly(Ephemeroptera Baetidae) from Utah USArsquo Entomological News 115 139ndash145

Wuillot J and Gillies MT (1993) lsquoCheleocloeon a new genus of Baetidae (Ephemeroptera)from West Africarsquo Revue drsquoHydrobiologie tropicale 26 213ndash217

Wuillot J and Gillies MT (1994) lsquoDicentroptilum a new genus of Mayflies (BaetidaeEphemeroptera) from Africarsquo Aquatic Insects 16 133ndash140

Zhou CF and Braasch D (2003) lsquoEine neue Gattung und Art der Heptageniidae ausdem ostlichen China (Ephemeroptera)rsquo Entomologische Nachrichten und Berichte 47147ndash151

Zhou CF Gui H and Su CR (2000) lsquoThe first record of the genus Brachycercus in Chinawith description of a new species (Ephemeroptera Caenidae)rsquo Entomologia Sinica 7132ndash134

Zhou CF and Peters JG (2003) lsquoThe nymph of Siphluriscus chinensis and additionalimaginal description A living mayfly with Jurassic origins (Siphluriscidae new familyEphemeroptera)rsquo Florida Entomologist 86 345ndash352

Zhou CF and Zheng LY (2001) lsquoRhithrogena trispina sp n a new species from China(Ephemeroptera Heptageniidae)rsquo Aquatic Insects 23 323ndash326

Zhou CF and Zheng LY (2003) lsquoTwo synonyms and one new species of the genusEphemera from China (Ephemeroptera Ephemeridae)rsquo Acta Zootaxonomica Sinica 28665ndash668

Zhou CF and Zheng LY (2004) lsquoThe genus Prosopistoma from China with description oftwo new species (Ephemeroptera Prosopistomatidae)rsquo Aquatic Insects 26 3ndash8

62 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Gattolliat J-L and Sartori M (2003) lsquoAn overview of the Baetidae of Madagascarrsquo inResearch Update on Ephemeroptera and Plecoptera ed E Gaino Perugia University ofPerugia Italy pp 135ndash144

Gattolliat J-L and Sartori M (2008) lsquoOrder Ephemeropterarsquo in Arthropod Fauna of theUAE ed A van Harten Abu Dhabi Dar Al Ummah pp 47ndash83

Gillies MT (1949) lsquoNotes on some Ephemeroptera Baetidae from India and South-EastAsiarsquo Transactions of the Royal Entomological Society of London 100 161ndash177

Gillies MT (1985) lsquoA preliminary account of the East African species of Cloeon Leach andRhithrocloeon gen n (Ephemeroptera)rsquo Aquatic Insects 7 1ndash17

Gillies MT (1988) lsquoDescription of the nymphs of some Afrotropical Baetidae (Ephemer-optera) I Cloeon Leach and Rhithrocloeon Gilliesrsquo Aquatic Insects 10 49ndash59

Gillies MT (1990) lsquoA revision of the African species of Centroptilum Eaton (BaetidaeEphemeroptera)rsquo Aquatic Insects 12 97ndash128

Gillies MT (1991a) lsquoA diphyletic origin for the two-tailed baetid nymphs occurring inEast African stony streams with a description of the new genus and species Tanzaniellaspinosa gen sp novrsquo in Overview and strategies of Ephemeroptera and Plecoptera edsJ Alba-Tercedor and A Sanchez-Ortega Gainesville FL Sandhill Crane Press Incpp 175ndash187

Gillies MT (1991b) lsquoA new species of Afroptilum (Afroptiloides) from East Africa (EphemBaetidae)rsquo Entomologistrsquos Monthly Magazine 127 109ndash115

Gillies MT (1994) lsquoDescriptions of some Afrotropical Baetidae (Ephemeroptera) II BaetisLeach sl East African speciesrsquo Aquatic Insects 16 105ndash118

Gillies MT (1998) lsquoThe mayflies (Ephemeroptera) of the upper River Sigi North-EastTanzaniarsquo Freshwater Forum 10 49ndash57

Gillies MT and Elouard J-M (1990) lsquoThe mayfly-mussel association a new example fromthe River Niger Basinrsquo in Mayflies and Stoneflies Life story and Biology ed ICCampbell Dordrecht Kluwer Academic Publishers pp 289ndash298

Gillies MT Elouard J-M and Wuillot J (1990) lsquoEphemeroptera from West Africa thegenus Ophelmatostoma (Baetidae)rsquo Revue drsquoHydrobiologie Tropicale 23(2) 115ndash120

Gillies MT and Wuillot J (1997) lsquoPlatycloeon a new genus of Baetidae (Ephemeroptera)from East Africarsquo Aquatic Insects 19 185ndash189

Glazaczow A (1997) lsquoObservations on the Psammophilous mayfly species Procleon nanumin the North East of Polandrsquo in Ephemeroptera amp Plecoptera Biology-Ecology-Systematics eds P Landolt and M Sartori Fribourg Mauron thorn Tinguely amp LachatSA pp 83ndash87

Gose K (1980) lsquoNihon san kagero-rui [The mayflies of Japanese Key to families genera andspecies]rsquo (in Japanese) Aquabiology (Nara) 2(1) 76ndash79 2(2) 122ndash123 2(3) 211ndash2152(4) 286ndash288 2(5) 366ndash368 2(6) 454ndash457

Hagen H (1858) lsquoSynopsis der Neuroptera Ceylonsrsquo Verhandlungen der Kaiserlich-Koniglichen Zoologisch-Botanischen Gesellschaft in Wien 8 471ndash488

Hagen H (1861) lsquoSynopsis of the Neuroptera of North America with a list of the SouthAmerican speciesrsquo Smithsonian Miscellaneous collections 1ndash347

Harker JE (1950) lsquoAustralian Ephemeroptera Part I Taxonomy of New South Walesspecies and evaluation of taxonomic charactersrsquo Proceedings of the Linnean Society ofNew South Wales 75 1ndash34

Harker JE (1954) lsquoThe Ephemeroptera from Eastern Australiarsquo Transactions of the RoyalEntomological Society of London 105 241ndash268

Harker JE (1957) lsquoSome new Australian Ephemeropterarsquo Proceedings of the RoyalEntomological Society of London B 26 63ndash78

Jacobus LM and McCafferty WP (2005a) lsquoApobaetis futilis (McDunnough) a newcombination in Nearctic Baetidae (Ephemeroptera)rsquo Proceedings of the EntomologicalSociety of Washington 107 979

Jacobus LM and McCafferty WP (2005b) lsquoA new species and new synonymin Camelobaetidius Demoulin (Ephemeroptera Baetidae)rsquo Journal of the KansasEntomological Society 78 153ndash157

Jacobus LM and McCafferty WP (2006) lsquoA new species of Acentrella Bengtsson(Ephemeroptera Baetidae) from Great Smoky Mountains National Park USArsquo AquaticInsects 28 101ndash111

56 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Kang SC Chang HC and Yang CT (1994) lsquoA revision of the genus Baetis in Taiwan(Ephemeroptera Baetidae)rsquo Journal of Taiwan Museum 47 9ndash44

Kang SC and Yang CT (1996) lsquoTwo new species of Baetis Leach (EphemeropteraBaetidae) from Taiwanrsquo Chinese Journal of Entomology 16 61ndash66

Kapur AP and Kripalani MB (1963) lsquoThe mayflies (Ephemeroptera) from the north-western Himalayarsquo Records of the Indian Museum 59 183ndash221 (1961)

Kazlauskas R (1963) lsquoNew and little known may flies (Ephemeroptera) from the USSRrsquoRevue drsquoEntomologie de lrsquoURSS 42 582ndash593

Kimmins DE (1947) lsquoNew species of Indian Ephemeropterarsquo Proceedings of the RoyalEntomological Society of London B 16 92ndash100

Kimmins DE (1956) lsquoNew species of Ephemeroptera from Ugandarsquo Bulletin of the BritishMuseum (Natural History) Entomology 4 71ndash87

Kopelke J-P (1980) lsquoEphemeroptera aus der Emergenz des zentralafrikanischen BergbachesKalengo (Zaıre) I Baetidaersquo Entomologische Abhandlungen 43 99ndash129

Kluge N Ju (2009) lsquoNew version of the database lsquolsquoEphemeroptera of the Worldrsquorsquo as the firstexperience of a permanent and objective web catalogue in biologyrsquo in InternationalPerspectives in Mayfly and Stonefly Research Proceedings of the 12th InternationalConference on Ephemeroptera and the 16th International Symposium on PlecopteraStuttgart 2008 ed AH Staniczek Aquatic Insects 31 Supplement 1 167ndash180

Lestage J-A (1918) lsquoLes Ephemeres drsquoAfrique (notes critiques sur les especes connues)rsquoRevue zoologique africaine 6 65ndash114

Leveque C Hougard JM Resh V Statzner B and Yameogo L (2003) lsquoFreshwaterecology and biodiversity in the tropics what did we learn from 30 years of onchocerciasiscontrol and the associated biomonitoring of West African riversrsquo Hydrobiologia 50023ndash49

Lugo-Ortiz CR (1999) lsquoSystematic studies of Baetidae (Insecta Ephemeroptera) withemphasis on biodiversity in the Southern Hemispherersquo Purdue University Department ofEntomology

Lugo-Ortiz CR and McCafferty WP (1995) lsquoThree distinctive new genera of Baetidae(Insecta Ephemeroptera) from South Americarsquo Annales de Limnologie 31 233ndash243

Lugo-Ortiz CR and McCafferty WP (1996a) lsquoAturbina georgei gen et sp n A smallminnow mayfly (Ephemeroptera Baetidae) without turbinate eyesrsquo Aquatic Insects 18175ndash183

Lugo-Ortiz CR and McCafferty WP (1996b) lsquoThe Bugilliesia complex of AfricanBaetidae (Ephemeroptera)rsquo Transactions of the American Entomological Society 122175ndash197

Lugo-Ortiz CR and McCafferty WP (1997) lsquoNew Afrotropical genus of Baetidae(Insecta Ephemeroptera) with bladelike mandiblesrsquo Bulletin de la Societe drsquoHistoireNaturelle de Toulouse 133 41ndash46

Lugo-Ortiz CR and McCafferty WP (1998a) lsquoThe Centroptiloides Complex ofAfrotropical small minnow mayflies (Ephemeroptera Baetidae)rsquo Annals of TheEntomological Society of America 91 1ndash26

Lugo-Ortiz CR and McCafferty WP (1998b) lsquoFirst report and new species of the genusCloeodes (Ephemeroptera Baetidae) from Australiarsquo Entomological News 109 122ndash128

Lugo-Ortiz CR and McCafferty WP (1998c) lsquoFive new genera of Baetidae (InsectaEphemeroptera) from South Americarsquo Annales de Limnologie 34 57ndash73

Lugo-Ortiz CR and McCafferty WP (1998d) lsquoA new North American genus ofBaetidae (Ephemeroptera) and key to Baetis complex generarsquo Entomological News 109345ndash353

Lugo-Ortiz CR and McCafferty WP (1998e) lsquoOffadens a new genus of small minnowmayflies (Ephemeroptera Baetidae) from Australiarsquo Proceedings of the EntomologicalSociety of Washington 100 306ndash309

Lugo-Ortiz CR and McCafferty WP (1998f) lsquoPhylogeny and biogeography of Nesydemiusn gen and related Afrotropical genera (Insecta Ephemeroptera Baetidae)rsquo Bulletin de laSociete drsquoHistoire Naturelle de Toulouse 134 7ndash12

Lugo-Ortiz CR and McCafferty WP (1999a) lsquoEdmundsiops instigatus A new genusand species of small minnow mayflies (Ephemeroptera Baetidae) from AustraliarsquoEntomological News 110 65ndash69

Aquatic Insects 57

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Lugo-Ortiz CR and McCafferty WP (1999b) lsquoGlobal biodiversity of the mayfly familyBaetidae (Ephemeroptera) a generic perspectiversquo Trends in Entomology 2 45ndash54

Lugo-Ortiz CR and McCafferty WP (1999c) lsquoA new genus of small minnow mayflies(Ephemeroptera Baetidae) with six new species from New Guinea and New BritainrsquoAnnales de Limnologie 35 57ndash70

Lugo-Ortiz CR and McCafferty WP (1999d) lsquoRevision of South American species ofBaetidae (Ephemeroptera) previously placed in Baetis Leach and Pseudocloeon KlapalekrsquoAnnales de Limnologie 35 257ndash262

Lugo-Ortiz CR and McCafferty WP (1999e) lsquoThree new genera of small minnow mayflies(Ephemeroptera Baetidae) from the Andes and Patagoniarsquo Studies on Neotropical Faunaand Environment 34 88ndash104

Lugo-Ortiz CR McCafferty WP and Waltz RD (1999) lsquoDefinition and reorganizationof the genus Pseudocloeon (Ephemeroptera Baetidae) with new species descriptions andcombinationsrsquo Transactions of the American Entomological Society 125 1ndash37

Mayo VK (1973) lsquoFour news species of the genus Baetodes (Ephemeroptera Baetidae)rsquo ThePan-Pacific Entomologist 49 308ndash314

McCafferty WP (1998) lsquoEphemeroptera and the great American interchangersquo Journal of theNorth American Benthological Society 17 1ndash20

McCafferty WP (2000) lsquoNotations on South American Baetidae (Ephemeroptera)rsquoEntomological News 111 375ndash379

McCafferty WP (2009) lsquoMayfly Centralrsquo wwwentmpurdueeduEntomologyresearchmayflymayflyhtml

McCafferty WP and Baumgardner DE (2003) lsquoLugoiops maya a new genus and species ofEphemeroptera (Baetidae) from Central Americarsquo Proceedings of the EntomologicalSociety of Washington 105 397ndash406

McCafferty WP and Sun L (2005) lsquoMystaxiops A new genus of small minnow mayflies(Ephemeroptera Baetidae) from Papua New Guinearsquo Proceedings of the EntomologicalSociety of Washington 107 536ndash542

McDunnough J (1922) lsquoTwo new Canadian May-flies (Ephemeridae)rsquo Canadian Entomol-ogist 53 117ndash120

McDunnough J (1923) lsquoNew Canadian Ephemeridae with notesrsquo Canadian Entomologist55 39ndash50

McDunnough J (1924) lsquoNew Ephemeridae from Illinoisrsquo Canadian Entomologist 56 7ndash9McDunnough J (1925) lsquoNew Canadian Ephemeridae with notes IIIrsquo Canadian Entomol-

ogist 57(168ndash176) 185ndash192McDunnough J (1926) lsquoNew Canadian Ephemeridae with notes IVrsquo Canadian Entomol-

ogist 58 296ndash303McDunnough J (1929) lsquoNotes on North American Ephemeroptera with descriptions of new

species IIrsquo Canadian Entomologist 61 169ndash180McDunnough J (1931) lsquoNew species of North American Ephemeropterarsquo Canadian

Entomologist 63 82ndash93McDunnough J (1932) lsquoNew species of North American Ephemeroptera IIrsquo Canadian

Entomologist 64 209ndash215McDunnough J (1936) lsquoA new Arctic baetid (Ephemeroptera)rsquo Canadian Entomologist 68

32ndash34McDunnough J (1939) lsquoNew British Columbian Ephemeropterarsquo Canadian Entomologist

71 49ndash54Mol AWM (1986) lsquoHarpagobaetis gulosus gen nov sp nov a new mayfly from Suriname

(Ephemeroptera Baetidae)rsquo Zoologische Mededelingen 60 63ndash70Mol AWM (1989) lsquoEchinobaetis phagas gen nov spec nov a new mayfly from Sulawesi

(Ephemeroptera Baetidae)rsquo Zoologische Mededelingen 63 61ndash72Monaghan MT Gattolliat JL Sartori M Elouard JM James H Derleth P Glaizot

O de Moor F and Vogler AP (2005) lsquoTrans-oceanic and endemic origins of the smallminnow mayflies (Ephemeroptera Baetidae) of Madagascarrsquo Proceedings of The RoyalSociety B-Biological Sciences 272 1829ndash1836

Muller-Liebenau I (1980a) lsquoJubabaetis gen n and Platybaetis gen n two new genera of thefamily Baetidae from the Oriental Regionrsquo in Advances in Ephemeroptera Biology edsJF Flannagan and KE Marshall New York Plenum Press pp 103ndash114

58 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Muller-Liebenau I (1980b) lsquoA new species of the genus Platybaetis Muller-Liebenau 1980P bishopi spn from Malaysia (Insecta Ephemeroptera)rsquo Gewasser und Abwasser 666795ndash101

Muller-Liebenau I (1981) lsquoReview of the original material of the baetid genera Baetis andPseudocloeon from the Sunda Islands and the Philippines described by G Ulmer withsome general remarks (Insecta Ephemeroptera)rsquo Mitteilungen der hamburger zoologischerMuseum und Institut 78 197ndash208

Muller-Liebenau I (1982a) lsquoFive new species of Pseudocloeon Klapalek 1905 (FamBaetidae) from the Oriental region (Insecta Ephemeroptera) with some general remarkson Pseudocloeonrsquo Archiv fur Hydrobiologie 95 283ndash298

Muller-Liebenau I (1982b) lsquoA new genus and species of Baetidae from Sri Lanka (Ceylon)Indocloeon primum gen n sp n (Insecta Ephemeroptera)rsquo Aquatic Insects 4 125ndash129

Muller-Liebenau I (1982c) lsquoNew species of the family Baetidae from the Philippines (InsectaEphemeroptera)rsquo Archiv fur Hydrobiologie 94 70ndash82

Muller-Liebenau I (1983) lsquoThree new species of the genus Centroptella Braasch ampSoldan 1980 from Sri Lanka (Insecta Ephemeroptera)rsquo Archiv fur Hydrobiologie 97486ndash500

Muller-Liebenau I (1984a) lsquoBaetidae from Sabah (East Malaysia) (Ephemeroptera)rsquo inProceeding of the fourth international conference on Ephemeroptera eds V LandaT Soldan and M Tonner Bechyne CSAV pp 85ndash99

Muller-Liebenau I (1984b) lsquoNew genera and species of the family Baetidae from West-Malaysia (River Gombak) (Insecta Ephemeroptera)rsquo Spixiana 7 253ndash284

Muller-Liebenau I (1985) lsquoBaetidae from Taiwan with remarks on Baetiella Ueno 1931(Insecta Ephemeroptera)rsquo Archiv fur Hydrobiologie 104 93ndash110

Muller-Liebenau I and Heard WH (1979) lsquoSymbiocloeon a new genus of Baetidaefrom Thailand (Insecta Ephemeroptera)rsquo in Proceedings of the Second InternationalConference on Ephemeroptera eds K Pasternak and R Sowa Warszawa PanstwoweWydawnictwo Naukowe pp 57ndash66

Muller-Liebenau I and Hubbard MD (1985) lsquoBaetidae from Sri Lanka with some generalremarks on the Baetidae of the Oriental region (Insecta Ephemeroptera)rsquo The FloridaEntomologist 68 537ndash561

Muller-Liebenau I and Morihara DK (1982) lsquoIndobaetis A new genus of Baetidae fromSri Lanka (Insecta Ephemeroptera) with two new speciesrsquo Gewasser und Abwasser 686926ndash34

Navas L (1911) lsquoNotas entomologicas Excursiones por los alrededores de Granadarsquo Boletinde la Sociedad Aragonesa de Ciencias Naturales 10 204ndash211

Navas L (1913) lsquoNotas entomologicas Excursiones por los alrededores de Zaragozarsquo Boletinde la Sociedad Aragonesa de Ciencias Naturales 12 75ndash77

Navas L (1917) lsquoNeuropteros nuevos o poco concocidoslsquo Memorias de la Real Academia deCiencias y Artes de Barcelona 13 393ndash395

Navas L (1930) lsquoInsectes du Congo Belge (Serie IV)rsquo Revue de Zoologie et de BotaniqueAfricaines 19 305ndash336

Navas L (1931a) lsquoInsectes du Congo Belge (Serie V)rsquo Revue de Zoologie et de BotaniqueAfricaines 20 257ndash279

Navas L (1931b) lsquoNevropteres et insectes voisins - Chine et pays environnants - 2e SeriersquoNotes drsquoentomologie chinoise 1 1ndash10

Needham JG Traver JR and Hsu Y-S (1935) The biology of mayflies with a systematicaccount of North American species Ithaca NY Comstock Publishing Company

Nieto C (2003a) lsquoEl genero Camelobaetidius (Ephemeroptera Baetidae) en la ArgentinarsquoActa Zoologica Mexicana 88 233ndash255

Nieto C (2003b) lsquoA new species of Guajirolus (Ephemeroptera Baetidae) from Argentinaand description of a new genus from Boliviarsquo in Research Update on Ephemeroptera andPlecoptera ed E Gaino Perugia University of Perugia Italy pp 153ndash158

Nieto C (2004a) lsquoThe genus Baetodes (Ephemeroptera Baetidae) in South America with thedescription of new species from Argentina Bolivia and Perursquo Studies on NeotropicalFauna and Environment 39 63ndash79

Nieto C (2004b) lsquoRedescription of Varipes (Ephemeroptera Baetidae) with the descriptionof new species from Bolivia and Argentinarsquo Aquatic Insects 26 161ndash173

Aquatic Insects 59

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Nieto C (2004c) lsquoSouth American Baetidae (Ephemeroptera) a new generic synonymyrsquoStudies on Neotropical Fauna and Environment 39 95ndash101

Nieto C and Richard B (2008) lsquoThe genus Cloeodes (Ephemeroptera Baetidae) inArgentina with new generic synonymy and new speciesrsquo Zootaxa 1727 1ndash21

Nieto C and Salles FF (2006) lsquoRevision of the genus Paracloeodes (EphemeropteraBaetidae) in South Americarsquo Zootaxa 1ndash33

Nolte U Tietbohl RS and McCafferty WP (1996) lsquoA mayfly from tropical Brazilcapable of tolerating short-term dehydrationrsquo Journal of the North American BenthologicalSociety 15 87ndash94

Park SY Bae YJ and Yoon IB (1996) lsquoRevision of the Baetidae (Ephemeroptera) ofKorea (1) Historical review Acentrella Bengtsson and Baetiella Uenorsquo EntomologicalResearch Bulletin 22 55ndash66

Peters WL (2001) lsquoThe Ephemeroptera of New Caledonia and New Zealandrsquo in Trends inResearch in Ephemeroptera and Plecoptera ed E Domınguez New York KluwerAcademicPlenum Publishers pp 43ndash45

Pictet F-J (1843ndash1845) Histoire naturelle des Insectes Nevropteres Famille des EphemerinesGeneve

Rambur P (1842) Histoire Naturelle des Insectes Nevropteres Paris Librairie Encyclope-dique de Roret

Salles FF (2007) lsquoThe presence of Chane Nieto and Guajirolus Flowers (EphemeropteraBaetidae) in Brazil with the description of a new speciesrsquo Revista Brasileira DeEntomologia 51 404ndash409

Salles FF Andrade MB and Da-Silva ER (2005) lsquoCamelobaetidius francischettii a newspecies of Baetidae (Ephemeroptera) from Brazilrsquo Zootaxa 47ndash53

Salles FF and Batista JD (2004) lsquoThe presence of Varipes Lugo-Ortiz amp McCafferty(Ephemeroptera Baetidae) in Brazil with the description of a new speciesrsquo Zootaxa 4561ndash6

Salles FF Batista JD and Soares Cabette HR (2004) lsquoBaetidae (Insecta Ephemer-optera) de Nova Xavantina Mato Grosso Brasil novos registros e descricao de una novaespecie de Cloeodes Traverrsquo Biota Neotropica 4 1ndash8

Salles FF and Francischetti CN (2004) lsquoCryptonympha dasilvai sp nov (EphemeropteraBaetidae) do Brasilrsquo Neotropical Entomology 33 213ndash216

Salles FF and Polegatto CM (2008) lsquoTwo new species of Baetodes Needham amp Murphy(Ephemeroptera Baetidae) from Brazilrsquo Zootaxa 43ndash50

Sartori M Derleth P and Gattolliat JL (2003) lsquoNew data about the mayflies(Ephemeroptera) from Borneorsquo in Research Update on Ephemeroptera and Plecopteraed E Gaino Perugia University of Perugia Italy pp 403ndash406

Soldan T and Godunko RJ (2006) lsquoBaetis atlanticus n sp a new species of the subgenusRhodobaetis Jacob 2003 from Madeira Portugal (Ephemeroptera Baetidae)rsquo Genus 175ndash17

Soldan T and Thomas AGB (1983a) lsquoBaetis numidicus nsp Ephemeroptere nouveaudrsquoAlgerie (Baetidae)rsquo Annales de Limnologie 19 207ndash211

Soldan T and Thomas AGB (1983b) lsquoNew and little-known species of mayflies(Ephemeroptera) from Algeriarsquo Acta Entomologica Bohemoslovaca 80 356ndash376

Soldan T and Thomas AGB (1985) lsquoCentroptilum dimorphicum sp n a new species ofmayfly (Ephemeroptera Baetidae) from Algeriarsquo Acta Entomologica Bohemoslovaca 82180ndash186

Suter PJ (1979) lsquoA revised key to the Australian genera of mature mayfly(Ephemeroptera) nymphsrsquo Transactions of the Royal Society of South Australia 103 79ndash83

Suter PJ (1986) lsquoThe Ephemeroptera (Mayflies) of South Australiarsquo Records of the SouthAustralian Museum 19 339ndash397

Suter PJ (1997) lsquoPreliminary guide to the identification of nymphs of Australian Baetidmayflies (Insecta Ephemeroptera) found in flowing watersrsquo in Identification Guide No 14Albany Co-Operative Research Center for Freshwater Ecology

Suter PJ (2000) lsquoEdmundsiops hickmani spnov Offadens frater (Tillyard) nov comband description of the nymph of Cloeon tasmaniae Tillyard (EphemeropteraBaetidae) from Tasmaniarsquo Papers and Proceedings of the Royal Society of Tasmania134 63ndash73

60 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Suter PJ (2001) lsquoPlatybaetis gagadjuensis a new species from Northern Australia(Ephemeroptera Baetidae)rsquo in Trends in Research in Ephemeroptera and Plecopteraed E Dominguez New York Kluwer AcademicPlenum Publishers pp 359ndash364

Suter PJ and Pearson MJ (2001) lsquoRedescription of Bungona Harker with new synonymsin the Australian Baetidae (Insecta Ephemeroptera)rsquo Memoirs of the Museum of Victoria58 247ndash254

Thomas A and Gagneur J (1994) lsquoComplements et corrections a la faune desEphemeropteres drsquoAfrique du Nord 6 Alainites sadati n sp drsquoAlgerie (EphemeropteraBaetidae)rsquo Bulletin de la Societe drsquoHistoire Naturelle de Toulouse 130 43ndash45

Thomas A and Peru N (2003) lsquoLes Ephemeres de la Guyane Francaise 6 Description deWaltzoyphius roberti n sp (Ephemeroptera Baetidae)rsquo Bulletin de la Societe drsquoHistoireNaturelle de Toulouse 138 15ndash20 (2002)

Thomas AGB (1986) lsquoEphemeropteres du Sud-Ouest de la France IV Baetis catharus nspdes Pyrenees (Baetidae)rsquo Bulletin de la Societe drsquoHistoire Naturelle de Toulouse 122175ndash179

Thomas AGB (1998) lsquoA provisional checklist of the mayflies of North Africa(Ephemeroptera)rsquo Bulletin de la Societe drsquoHistoire Naturelle de Toulouse 134 13ndash20

Thomas AGB and Dia A (1984) lsquoBaetis baroukianus nsp Ephemeroptere nouveau duLiban (Baetidae)rsquo Bulletin de la Societe drsquoHistoire Naturelle de Toulouse 120 7ndash12

Thomas AGB and Dia A (1985) lsquoBaetis spei nsp Ephemeroptere nouveau du Libandescription et ecologie (Baetidae)rsquo Annales de Limnologie 21 241ndash245

Thomas AGB and Gazagnes G (1984) lsquoBaetis cyrneus n sp Ephemeroptere nouveau deCorse (Baetidae)rsquo Annales de Limnologie 20 199ndash202

Thomas AGB and Sartori M (1989) lsquoMayflies (Insecta Ephemeroptera) of SaudiArabiarsquo Fauna of Saudi Arabia 10 87ndash94

Thomas AGB and Soldan T (1987) lsquoBaetis ingridae n sp Ephemeroptere nouveau deCorse (Baetidae)rsquo Annales de Limnologie 23 23ndash26

Tillyard RJ (1936) lsquoThe trout-food insects of Tasmania Part II- A monograph of themayflies of Tasmaniarsquo Papers and proceedings of the Royal Society of Tasmania 193523ndash59

Tong X and Dudgeon D (2000) lsquoBaetiella (Ephemeroptera Baetidae) in Hong Kong withdescription of a new speciesrsquo Entomological News 111 143ndash148

Tong X and Dudgeon D (2003) lsquoFirst record of the genus Chopralla (EphemeropteraBaetidae) from China and description of a new speciesrsquo The Raffles Bulletin of Zoology51 17ndash19

Traver JR and Edmunds GF (1968) lsquoA revision of the Baetidae with spatulate-clawednymphs (Ephemeroptera)rsquo Pacific Insects 10 629ndash677

Ulmer G (1909) lsquoEphemeriden von Madagaskar und den Comorenrsquo Voeltzkowrsquos Reise inOstrafrika 1903ndash1905 II 365ndash368

Ulmer G (1939) lsquoEintagsfliegen (Ephemeropteren) von den Sunda-Inselnrsquo Archiv furHydrobiologie 16 443ndash692

Walsh BD (1863) lsquoObservations on certain N A Neuroptera by H Hagen M D ofKoenigsberg Prussia translated from the original French ms and published bypermission of the author with notes and descriptions of about twenty new N A speciesof Pseudoneuropterarsquo Proceedings of the Entomological Society of Philadelphia 2167ndash272

Walsh BD (1863) lsquoList of the Pseudoneuroptera of Illinois contained in the cabinet of thewriter with descriptions of over forty new species and notes on their structural affinitiesrsquoProceedings of the Academy of Natural Sciences of Philadelphia 1862 361ndash402

Waltz RD and McCafferty WP (1985) lsquoMoribaetis a new genus of NeotropicalBaetidae (Ephemeroptera)rsquo Proceedings of the Entomological Society of Washington 87239ndash251

Waltz RD and McCafferty WP (1986) lsquoApobaetis etowah (Traver) a new combination inNearctic Baetidae (Ephemeroptera)rsquo Proceedings of the Entomological Society ofWashington 88 191

Waltz RD and McCafferty WP (1987a) lsquoGeneric revision of Cloeodes and description oftwo new genera (Ephemeroptera Baetidae)rsquo Proceedings of the Entomological Society ofWashington 89 177ndash184

Aquatic Insects 61

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Waltz RD and McCafferty WP (1987b) lsquoNew genera of Baetidae for some Nearcticpreviously included in Baetis Leach (Ephemeroptera)rsquo Annals of the Entomological Societyof America 80 667ndash670

Waltz RD and McCafferty WP (1987c) lsquoSystematics of Pseudocloeon AcentrellaBaetiella and Liebebiella new genus (Ephemeroptera Baetidae)rsquo Journal of New YorkEntomological Society 95 553ndash568

Waltz RD and McCafferty WP (1989) lsquoNew species redescriptions and cladisticsof the genus Pseudocentroptiloides (Epherneroptera Baetidae)rsquo Journal of New YorkEntomological Society 97 151ndash158

Waltz RD McCafferty WP and Thomas A (1994) lsquoSystematics of Alainites n genDiphetor Indobaetis Nigrobaetis n stat and Takobia n stat (Ephemeroptera Baetidae)rsquoBulletin de la Societe drsquoHistoire Naturelle de Toulouse 130 33ndash36

Wang T-Q and McCafferty WP (1996) lsquoNew diagnostic characters for the mayfly familyBaetidae (Ephemeroptera)rsquo Entomological News 107 207ndash212

Weyenbergh H (1883) lsquoBijdrage tot de Kennis der zuid-amerikaansche EphemeridenrsquoTijdschrift voor Entomologie 26 159ndash174

Wiersema NA (2000) lsquoA new combination for two North American small minnow mayflies(Ephemeroptera Baetidae)rsquo Entomological News 111 140ndash142

Wiersema NA and McCafferty WP (2004) lsquoNew specific synonyms and records of NorthAmerican Centroptilum and Procloeon (Ephemeroptera Baetidae)rsquo Entomological News115 121ndash128

Wiersema NA Nelson CR and Kuehnl KF (2004) lsquoA new small minnow mayfly(Ephemeroptera Baetidae) from Utah USArsquo Entomological News 115 139ndash145

Wuillot J and Gillies MT (1993) lsquoCheleocloeon a new genus of Baetidae (Ephemeroptera)from West Africarsquo Revue drsquoHydrobiologie tropicale 26 213ndash217

Wuillot J and Gillies MT (1994) lsquoDicentroptilum a new genus of Mayflies (BaetidaeEphemeroptera) from Africarsquo Aquatic Insects 16 133ndash140

Zhou CF and Braasch D (2003) lsquoEine neue Gattung und Art der Heptageniidae ausdem ostlichen China (Ephemeroptera)rsquo Entomologische Nachrichten und Berichte 47147ndash151

Zhou CF Gui H and Su CR (2000) lsquoThe first record of the genus Brachycercus in Chinawith description of a new species (Ephemeroptera Caenidae)rsquo Entomologia Sinica 7132ndash134

Zhou CF and Peters JG (2003) lsquoThe nymph of Siphluriscus chinensis and additionalimaginal description A living mayfly with Jurassic origins (Siphluriscidae new familyEphemeroptera)rsquo Florida Entomologist 86 345ndash352

Zhou CF and Zheng LY (2001) lsquoRhithrogena trispina sp n a new species from China(Ephemeroptera Heptageniidae)rsquo Aquatic Insects 23 323ndash326

Zhou CF and Zheng LY (2003) lsquoTwo synonyms and one new species of the genusEphemera from China (Ephemeroptera Ephemeridae)rsquo Acta Zootaxonomica Sinica 28665ndash668

Zhou CF and Zheng LY (2004) lsquoThe genus Prosopistoma from China with description oftwo new species (Ephemeroptera Prosopistomatidae)rsquo Aquatic Insects 26 3ndash8

62 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Kang SC Chang HC and Yang CT (1994) lsquoA revision of the genus Baetis in Taiwan(Ephemeroptera Baetidae)rsquo Journal of Taiwan Museum 47 9ndash44

Kang SC and Yang CT (1996) lsquoTwo new species of Baetis Leach (EphemeropteraBaetidae) from Taiwanrsquo Chinese Journal of Entomology 16 61ndash66

Kapur AP and Kripalani MB (1963) lsquoThe mayflies (Ephemeroptera) from the north-western Himalayarsquo Records of the Indian Museum 59 183ndash221 (1961)

Kazlauskas R (1963) lsquoNew and little known may flies (Ephemeroptera) from the USSRrsquoRevue drsquoEntomologie de lrsquoURSS 42 582ndash593

Kimmins DE (1947) lsquoNew species of Indian Ephemeropterarsquo Proceedings of the RoyalEntomological Society of London B 16 92ndash100

Kimmins DE (1956) lsquoNew species of Ephemeroptera from Ugandarsquo Bulletin of the BritishMuseum (Natural History) Entomology 4 71ndash87

Kopelke J-P (1980) lsquoEphemeroptera aus der Emergenz des zentralafrikanischen BergbachesKalengo (Zaıre) I Baetidaersquo Entomologische Abhandlungen 43 99ndash129

Kluge N Ju (2009) lsquoNew version of the database lsquolsquoEphemeroptera of the Worldrsquorsquo as the firstexperience of a permanent and objective web catalogue in biologyrsquo in InternationalPerspectives in Mayfly and Stonefly Research Proceedings of the 12th InternationalConference on Ephemeroptera and the 16th International Symposium on PlecopteraStuttgart 2008 ed AH Staniczek Aquatic Insects 31 Supplement 1 167ndash180

Lestage J-A (1918) lsquoLes Ephemeres drsquoAfrique (notes critiques sur les especes connues)rsquoRevue zoologique africaine 6 65ndash114

Leveque C Hougard JM Resh V Statzner B and Yameogo L (2003) lsquoFreshwaterecology and biodiversity in the tropics what did we learn from 30 years of onchocerciasiscontrol and the associated biomonitoring of West African riversrsquo Hydrobiologia 50023ndash49

Lugo-Ortiz CR (1999) lsquoSystematic studies of Baetidae (Insecta Ephemeroptera) withemphasis on biodiversity in the Southern Hemispherersquo Purdue University Department ofEntomology

Lugo-Ortiz CR and McCafferty WP (1995) lsquoThree distinctive new genera of Baetidae(Insecta Ephemeroptera) from South Americarsquo Annales de Limnologie 31 233ndash243

Lugo-Ortiz CR and McCafferty WP (1996a) lsquoAturbina georgei gen et sp n A smallminnow mayfly (Ephemeroptera Baetidae) without turbinate eyesrsquo Aquatic Insects 18175ndash183

Lugo-Ortiz CR and McCafferty WP (1996b) lsquoThe Bugilliesia complex of AfricanBaetidae (Ephemeroptera)rsquo Transactions of the American Entomological Society 122175ndash197

Lugo-Ortiz CR and McCafferty WP (1997) lsquoNew Afrotropical genus of Baetidae(Insecta Ephemeroptera) with bladelike mandiblesrsquo Bulletin de la Societe drsquoHistoireNaturelle de Toulouse 133 41ndash46

Lugo-Ortiz CR and McCafferty WP (1998a) lsquoThe Centroptiloides Complex ofAfrotropical small minnow mayflies (Ephemeroptera Baetidae)rsquo Annals of TheEntomological Society of America 91 1ndash26

Lugo-Ortiz CR and McCafferty WP (1998b) lsquoFirst report and new species of the genusCloeodes (Ephemeroptera Baetidae) from Australiarsquo Entomological News 109 122ndash128

Lugo-Ortiz CR and McCafferty WP (1998c) lsquoFive new genera of Baetidae (InsectaEphemeroptera) from South Americarsquo Annales de Limnologie 34 57ndash73

Lugo-Ortiz CR and McCafferty WP (1998d) lsquoA new North American genus ofBaetidae (Ephemeroptera) and key to Baetis complex generarsquo Entomological News 109345ndash353

Lugo-Ortiz CR and McCafferty WP (1998e) lsquoOffadens a new genus of small minnowmayflies (Ephemeroptera Baetidae) from Australiarsquo Proceedings of the EntomologicalSociety of Washington 100 306ndash309

Lugo-Ortiz CR and McCafferty WP (1998f) lsquoPhylogeny and biogeography of Nesydemiusn gen and related Afrotropical genera (Insecta Ephemeroptera Baetidae)rsquo Bulletin de laSociete drsquoHistoire Naturelle de Toulouse 134 7ndash12

Lugo-Ortiz CR and McCafferty WP (1999a) lsquoEdmundsiops instigatus A new genusand species of small minnow mayflies (Ephemeroptera Baetidae) from AustraliarsquoEntomological News 110 65ndash69

Aquatic Insects 57

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Lugo-Ortiz CR and McCafferty WP (1999b) lsquoGlobal biodiversity of the mayfly familyBaetidae (Ephemeroptera) a generic perspectiversquo Trends in Entomology 2 45ndash54

Lugo-Ortiz CR and McCafferty WP (1999c) lsquoA new genus of small minnow mayflies(Ephemeroptera Baetidae) with six new species from New Guinea and New BritainrsquoAnnales de Limnologie 35 57ndash70

Lugo-Ortiz CR and McCafferty WP (1999d) lsquoRevision of South American species ofBaetidae (Ephemeroptera) previously placed in Baetis Leach and Pseudocloeon KlapalekrsquoAnnales de Limnologie 35 257ndash262

Lugo-Ortiz CR and McCafferty WP (1999e) lsquoThree new genera of small minnow mayflies(Ephemeroptera Baetidae) from the Andes and Patagoniarsquo Studies on Neotropical Faunaand Environment 34 88ndash104

Lugo-Ortiz CR McCafferty WP and Waltz RD (1999) lsquoDefinition and reorganizationof the genus Pseudocloeon (Ephemeroptera Baetidae) with new species descriptions andcombinationsrsquo Transactions of the American Entomological Society 125 1ndash37

Mayo VK (1973) lsquoFour news species of the genus Baetodes (Ephemeroptera Baetidae)rsquo ThePan-Pacific Entomologist 49 308ndash314

McCafferty WP (1998) lsquoEphemeroptera and the great American interchangersquo Journal of theNorth American Benthological Society 17 1ndash20

McCafferty WP (2000) lsquoNotations on South American Baetidae (Ephemeroptera)rsquoEntomological News 111 375ndash379

McCafferty WP (2009) lsquoMayfly Centralrsquo wwwentmpurdueeduEntomologyresearchmayflymayflyhtml

McCafferty WP and Baumgardner DE (2003) lsquoLugoiops maya a new genus and species ofEphemeroptera (Baetidae) from Central Americarsquo Proceedings of the EntomologicalSociety of Washington 105 397ndash406

McCafferty WP and Sun L (2005) lsquoMystaxiops A new genus of small minnow mayflies(Ephemeroptera Baetidae) from Papua New Guinearsquo Proceedings of the EntomologicalSociety of Washington 107 536ndash542

McDunnough J (1922) lsquoTwo new Canadian May-flies (Ephemeridae)rsquo Canadian Entomol-ogist 53 117ndash120

McDunnough J (1923) lsquoNew Canadian Ephemeridae with notesrsquo Canadian Entomologist55 39ndash50

McDunnough J (1924) lsquoNew Ephemeridae from Illinoisrsquo Canadian Entomologist 56 7ndash9McDunnough J (1925) lsquoNew Canadian Ephemeridae with notes IIIrsquo Canadian Entomol-

ogist 57(168ndash176) 185ndash192McDunnough J (1926) lsquoNew Canadian Ephemeridae with notes IVrsquo Canadian Entomol-

ogist 58 296ndash303McDunnough J (1929) lsquoNotes on North American Ephemeroptera with descriptions of new

species IIrsquo Canadian Entomologist 61 169ndash180McDunnough J (1931) lsquoNew species of North American Ephemeropterarsquo Canadian

Entomologist 63 82ndash93McDunnough J (1932) lsquoNew species of North American Ephemeroptera IIrsquo Canadian

Entomologist 64 209ndash215McDunnough J (1936) lsquoA new Arctic baetid (Ephemeroptera)rsquo Canadian Entomologist 68

32ndash34McDunnough J (1939) lsquoNew British Columbian Ephemeropterarsquo Canadian Entomologist

71 49ndash54Mol AWM (1986) lsquoHarpagobaetis gulosus gen nov sp nov a new mayfly from Suriname

(Ephemeroptera Baetidae)rsquo Zoologische Mededelingen 60 63ndash70Mol AWM (1989) lsquoEchinobaetis phagas gen nov spec nov a new mayfly from Sulawesi

(Ephemeroptera Baetidae)rsquo Zoologische Mededelingen 63 61ndash72Monaghan MT Gattolliat JL Sartori M Elouard JM James H Derleth P Glaizot

O de Moor F and Vogler AP (2005) lsquoTrans-oceanic and endemic origins of the smallminnow mayflies (Ephemeroptera Baetidae) of Madagascarrsquo Proceedings of The RoyalSociety B-Biological Sciences 272 1829ndash1836

Muller-Liebenau I (1980a) lsquoJubabaetis gen n and Platybaetis gen n two new genera of thefamily Baetidae from the Oriental Regionrsquo in Advances in Ephemeroptera Biology edsJF Flannagan and KE Marshall New York Plenum Press pp 103ndash114

58 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Muller-Liebenau I (1980b) lsquoA new species of the genus Platybaetis Muller-Liebenau 1980P bishopi spn from Malaysia (Insecta Ephemeroptera)rsquo Gewasser und Abwasser 666795ndash101

Muller-Liebenau I (1981) lsquoReview of the original material of the baetid genera Baetis andPseudocloeon from the Sunda Islands and the Philippines described by G Ulmer withsome general remarks (Insecta Ephemeroptera)rsquo Mitteilungen der hamburger zoologischerMuseum und Institut 78 197ndash208

Muller-Liebenau I (1982a) lsquoFive new species of Pseudocloeon Klapalek 1905 (FamBaetidae) from the Oriental region (Insecta Ephemeroptera) with some general remarkson Pseudocloeonrsquo Archiv fur Hydrobiologie 95 283ndash298

Muller-Liebenau I (1982b) lsquoA new genus and species of Baetidae from Sri Lanka (Ceylon)Indocloeon primum gen n sp n (Insecta Ephemeroptera)rsquo Aquatic Insects 4 125ndash129

Muller-Liebenau I (1982c) lsquoNew species of the family Baetidae from the Philippines (InsectaEphemeroptera)rsquo Archiv fur Hydrobiologie 94 70ndash82

Muller-Liebenau I (1983) lsquoThree new species of the genus Centroptella Braasch ampSoldan 1980 from Sri Lanka (Insecta Ephemeroptera)rsquo Archiv fur Hydrobiologie 97486ndash500

Muller-Liebenau I (1984a) lsquoBaetidae from Sabah (East Malaysia) (Ephemeroptera)rsquo inProceeding of the fourth international conference on Ephemeroptera eds V LandaT Soldan and M Tonner Bechyne CSAV pp 85ndash99

Muller-Liebenau I (1984b) lsquoNew genera and species of the family Baetidae from West-Malaysia (River Gombak) (Insecta Ephemeroptera)rsquo Spixiana 7 253ndash284

Muller-Liebenau I (1985) lsquoBaetidae from Taiwan with remarks on Baetiella Ueno 1931(Insecta Ephemeroptera)rsquo Archiv fur Hydrobiologie 104 93ndash110

Muller-Liebenau I and Heard WH (1979) lsquoSymbiocloeon a new genus of Baetidaefrom Thailand (Insecta Ephemeroptera)rsquo in Proceedings of the Second InternationalConference on Ephemeroptera eds K Pasternak and R Sowa Warszawa PanstwoweWydawnictwo Naukowe pp 57ndash66

Muller-Liebenau I and Hubbard MD (1985) lsquoBaetidae from Sri Lanka with some generalremarks on the Baetidae of the Oriental region (Insecta Ephemeroptera)rsquo The FloridaEntomologist 68 537ndash561

Muller-Liebenau I and Morihara DK (1982) lsquoIndobaetis A new genus of Baetidae fromSri Lanka (Insecta Ephemeroptera) with two new speciesrsquo Gewasser und Abwasser 686926ndash34

Navas L (1911) lsquoNotas entomologicas Excursiones por los alrededores de Granadarsquo Boletinde la Sociedad Aragonesa de Ciencias Naturales 10 204ndash211

Navas L (1913) lsquoNotas entomologicas Excursiones por los alrededores de Zaragozarsquo Boletinde la Sociedad Aragonesa de Ciencias Naturales 12 75ndash77

Navas L (1917) lsquoNeuropteros nuevos o poco concocidoslsquo Memorias de la Real Academia deCiencias y Artes de Barcelona 13 393ndash395

Navas L (1930) lsquoInsectes du Congo Belge (Serie IV)rsquo Revue de Zoologie et de BotaniqueAfricaines 19 305ndash336

Navas L (1931a) lsquoInsectes du Congo Belge (Serie V)rsquo Revue de Zoologie et de BotaniqueAfricaines 20 257ndash279

Navas L (1931b) lsquoNevropteres et insectes voisins - Chine et pays environnants - 2e SeriersquoNotes drsquoentomologie chinoise 1 1ndash10

Needham JG Traver JR and Hsu Y-S (1935) The biology of mayflies with a systematicaccount of North American species Ithaca NY Comstock Publishing Company

Nieto C (2003a) lsquoEl genero Camelobaetidius (Ephemeroptera Baetidae) en la ArgentinarsquoActa Zoologica Mexicana 88 233ndash255

Nieto C (2003b) lsquoA new species of Guajirolus (Ephemeroptera Baetidae) from Argentinaand description of a new genus from Boliviarsquo in Research Update on Ephemeroptera andPlecoptera ed E Gaino Perugia University of Perugia Italy pp 153ndash158

Nieto C (2004a) lsquoThe genus Baetodes (Ephemeroptera Baetidae) in South America with thedescription of new species from Argentina Bolivia and Perursquo Studies on NeotropicalFauna and Environment 39 63ndash79

Nieto C (2004b) lsquoRedescription of Varipes (Ephemeroptera Baetidae) with the descriptionof new species from Bolivia and Argentinarsquo Aquatic Insects 26 161ndash173

Aquatic Insects 59

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Nieto C (2004c) lsquoSouth American Baetidae (Ephemeroptera) a new generic synonymyrsquoStudies on Neotropical Fauna and Environment 39 95ndash101

Nieto C and Richard B (2008) lsquoThe genus Cloeodes (Ephemeroptera Baetidae) inArgentina with new generic synonymy and new speciesrsquo Zootaxa 1727 1ndash21

Nieto C and Salles FF (2006) lsquoRevision of the genus Paracloeodes (EphemeropteraBaetidae) in South Americarsquo Zootaxa 1ndash33

Nolte U Tietbohl RS and McCafferty WP (1996) lsquoA mayfly from tropical Brazilcapable of tolerating short-term dehydrationrsquo Journal of the North American BenthologicalSociety 15 87ndash94

Park SY Bae YJ and Yoon IB (1996) lsquoRevision of the Baetidae (Ephemeroptera) ofKorea (1) Historical review Acentrella Bengtsson and Baetiella Uenorsquo EntomologicalResearch Bulletin 22 55ndash66

Peters WL (2001) lsquoThe Ephemeroptera of New Caledonia and New Zealandrsquo in Trends inResearch in Ephemeroptera and Plecoptera ed E Domınguez New York KluwerAcademicPlenum Publishers pp 43ndash45

Pictet F-J (1843ndash1845) Histoire naturelle des Insectes Nevropteres Famille des EphemerinesGeneve

Rambur P (1842) Histoire Naturelle des Insectes Nevropteres Paris Librairie Encyclope-dique de Roret

Salles FF (2007) lsquoThe presence of Chane Nieto and Guajirolus Flowers (EphemeropteraBaetidae) in Brazil with the description of a new speciesrsquo Revista Brasileira DeEntomologia 51 404ndash409

Salles FF Andrade MB and Da-Silva ER (2005) lsquoCamelobaetidius francischettii a newspecies of Baetidae (Ephemeroptera) from Brazilrsquo Zootaxa 47ndash53

Salles FF and Batista JD (2004) lsquoThe presence of Varipes Lugo-Ortiz amp McCafferty(Ephemeroptera Baetidae) in Brazil with the description of a new speciesrsquo Zootaxa 4561ndash6

Salles FF Batista JD and Soares Cabette HR (2004) lsquoBaetidae (Insecta Ephemer-optera) de Nova Xavantina Mato Grosso Brasil novos registros e descricao de una novaespecie de Cloeodes Traverrsquo Biota Neotropica 4 1ndash8

Salles FF and Francischetti CN (2004) lsquoCryptonympha dasilvai sp nov (EphemeropteraBaetidae) do Brasilrsquo Neotropical Entomology 33 213ndash216

Salles FF and Polegatto CM (2008) lsquoTwo new species of Baetodes Needham amp Murphy(Ephemeroptera Baetidae) from Brazilrsquo Zootaxa 43ndash50

Sartori M Derleth P and Gattolliat JL (2003) lsquoNew data about the mayflies(Ephemeroptera) from Borneorsquo in Research Update on Ephemeroptera and Plecopteraed E Gaino Perugia University of Perugia Italy pp 403ndash406

Soldan T and Godunko RJ (2006) lsquoBaetis atlanticus n sp a new species of the subgenusRhodobaetis Jacob 2003 from Madeira Portugal (Ephemeroptera Baetidae)rsquo Genus 175ndash17

Soldan T and Thomas AGB (1983a) lsquoBaetis numidicus nsp Ephemeroptere nouveaudrsquoAlgerie (Baetidae)rsquo Annales de Limnologie 19 207ndash211

Soldan T and Thomas AGB (1983b) lsquoNew and little-known species of mayflies(Ephemeroptera) from Algeriarsquo Acta Entomologica Bohemoslovaca 80 356ndash376

Soldan T and Thomas AGB (1985) lsquoCentroptilum dimorphicum sp n a new species ofmayfly (Ephemeroptera Baetidae) from Algeriarsquo Acta Entomologica Bohemoslovaca 82180ndash186

Suter PJ (1979) lsquoA revised key to the Australian genera of mature mayfly(Ephemeroptera) nymphsrsquo Transactions of the Royal Society of South Australia 103 79ndash83

Suter PJ (1986) lsquoThe Ephemeroptera (Mayflies) of South Australiarsquo Records of the SouthAustralian Museum 19 339ndash397

Suter PJ (1997) lsquoPreliminary guide to the identification of nymphs of Australian Baetidmayflies (Insecta Ephemeroptera) found in flowing watersrsquo in Identification Guide No 14Albany Co-Operative Research Center for Freshwater Ecology

Suter PJ (2000) lsquoEdmundsiops hickmani spnov Offadens frater (Tillyard) nov comband description of the nymph of Cloeon tasmaniae Tillyard (EphemeropteraBaetidae) from Tasmaniarsquo Papers and Proceedings of the Royal Society of Tasmania134 63ndash73

60 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Suter PJ (2001) lsquoPlatybaetis gagadjuensis a new species from Northern Australia(Ephemeroptera Baetidae)rsquo in Trends in Research in Ephemeroptera and Plecopteraed E Dominguez New York Kluwer AcademicPlenum Publishers pp 359ndash364

Suter PJ and Pearson MJ (2001) lsquoRedescription of Bungona Harker with new synonymsin the Australian Baetidae (Insecta Ephemeroptera)rsquo Memoirs of the Museum of Victoria58 247ndash254

Thomas A and Gagneur J (1994) lsquoComplements et corrections a la faune desEphemeropteres drsquoAfrique du Nord 6 Alainites sadati n sp drsquoAlgerie (EphemeropteraBaetidae)rsquo Bulletin de la Societe drsquoHistoire Naturelle de Toulouse 130 43ndash45

Thomas A and Peru N (2003) lsquoLes Ephemeres de la Guyane Francaise 6 Description deWaltzoyphius roberti n sp (Ephemeroptera Baetidae)rsquo Bulletin de la Societe drsquoHistoireNaturelle de Toulouse 138 15ndash20 (2002)

Thomas AGB (1986) lsquoEphemeropteres du Sud-Ouest de la France IV Baetis catharus nspdes Pyrenees (Baetidae)rsquo Bulletin de la Societe drsquoHistoire Naturelle de Toulouse 122175ndash179

Thomas AGB (1998) lsquoA provisional checklist of the mayflies of North Africa(Ephemeroptera)rsquo Bulletin de la Societe drsquoHistoire Naturelle de Toulouse 134 13ndash20

Thomas AGB and Dia A (1984) lsquoBaetis baroukianus nsp Ephemeroptere nouveau duLiban (Baetidae)rsquo Bulletin de la Societe drsquoHistoire Naturelle de Toulouse 120 7ndash12

Thomas AGB and Dia A (1985) lsquoBaetis spei nsp Ephemeroptere nouveau du Libandescription et ecologie (Baetidae)rsquo Annales de Limnologie 21 241ndash245

Thomas AGB and Gazagnes G (1984) lsquoBaetis cyrneus n sp Ephemeroptere nouveau deCorse (Baetidae)rsquo Annales de Limnologie 20 199ndash202

Thomas AGB and Sartori M (1989) lsquoMayflies (Insecta Ephemeroptera) of SaudiArabiarsquo Fauna of Saudi Arabia 10 87ndash94

Thomas AGB and Soldan T (1987) lsquoBaetis ingridae n sp Ephemeroptere nouveau deCorse (Baetidae)rsquo Annales de Limnologie 23 23ndash26

Tillyard RJ (1936) lsquoThe trout-food insects of Tasmania Part II- A monograph of themayflies of Tasmaniarsquo Papers and proceedings of the Royal Society of Tasmania 193523ndash59

Tong X and Dudgeon D (2000) lsquoBaetiella (Ephemeroptera Baetidae) in Hong Kong withdescription of a new speciesrsquo Entomological News 111 143ndash148

Tong X and Dudgeon D (2003) lsquoFirst record of the genus Chopralla (EphemeropteraBaetidae) from China and description of a new speciesrsquo The Raffles Bulletin of Zoology51 17ndash19

Traver JR and Edmunds GF (1968) lsquoA revision of the Baetidae with spatulate-clawednymphs (Ephemeroptera)rsquo Pacific Insects 10 629ndash677

Ulmer G (1909) lsquoEphemeriden von Madagaskar und den Comorenrsquo Voeltzkowrsquos Reise inOstrafrika 1903ndash1905 II 365ndash368

Ulmer G (1939) lsquoEintagsfliegen (Ephemeropteren) von den Sunda-Inselnrsquo Archiv furHydrobiologie 16 443ndash692

Walsh BD (1863) lsquoObservations on certain N A Neuroptera by H Hagen M D ofKoenigsberg Prussia translated from the original French ms and published bypermission of the author with notes and descriptions of about twenty new N A speciesof Pseudoneuropterarsquo Proceedings of the Entomological Society of Philadelphia 2167ndash272

Walsh BD (1863) lsquoList of the Pseudoneuroptera of Illinois contained in the cabinet of thewriter with descriptions of over forty new species and notes on their structural affinitiesrsquoProceedings of the Academy of Natural Sciences of Philadelphia 1862 361ndash402

Waltz RD and McCafferty WP (1985) lsquoMoribaetis a new genus of NeotropicalBaetidae (Ephemeroptera)rsquo Proceedings of the Entomological Society of Washington 87239ndash251

Waltz RD and McCafferty WP (1986) lsquoApobaetis etowah (Traver) a new combination inNearctic Baetidae (Ephemeroptera)rsquo Proceedings of the Entomological Society ofWashington 88 191

Waltz RD and McCafferty WP (1987a) lsquoGeneric revision of Cloeodes and description oftwo new genera (Ephemeroptera Baetidae)rsquo Proceedings of the Entomological Society ofWashington 89 177ndash184

Aquatic Insects 61

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Waltz RD and McCafferty WP (1987b) lsquoNew genera of Baetidae for some Nearcticpreviously included in Baetis Leach (Ephemeroptera)rsquo Annals of the Entomological Societyof America 80 667ndash670

Waltz RD and McCafferty WP (1987c) lsquoSystematics of Pseudocloeon AcentrellaBaetiella and Liebebiella new genus (Ephemeroptera Baetidae)rsquo Journal of New YorkEntomological Society 95 553ndash568

Waltz RD and McCafferty WP (1989) lsquoNew species redescriptions and cladisticsof the genus Pseudocentroptiloides (Epherneroptera Baetidae)rsquo Journal of New YorkEntomological Society 97 151ndash158

Waltz RD McCafferty WP and Thomas A (1994) lsquoSystematics of Alainites n genDiphetor Indobaetis Nigrobaetis n stat and Takobia n stat (Ephemeroptera Baetidae)rsquoBulletin de la Societe drsquoHistoire Naturelle de Toulouse 130 33ndash36

Wang T-Q and McCafferty WP (1996) lsquoNew diagnostic characters for the mayfly familyBaetidae (Ephemeroptera)rsquo Entomological News 107 207ndash212

Weyenbergh H (1883) lsquoBijdrage tot de Kennis der zuid-amerikaansche EphemeridenrsquoTijdschrift voor Entomologie 26 159ndash174

Wiersema NA (2000) lsquoA new combination for two North American small minnow mayflies(Ephemeroptera Baetidae)rsquo Entomological News 111 140ndash142

Wiersema NA and McCafferty WP (2004) lsquoNew specific synonyms and records of NorthAmerican Centroptilum and Procloeon (Ephemeroptera Baetidae)rsquo Entomological News115 121ndash128

Wiersema NA Nelson CR and Kuehnl KF (2004) lsquoA new small minnow mayfly(Ephemeroptera Baetidae) from Utah USArsquo Entomological News 115 139ndash145

Wuillot J and Gillies MT (1993) lsquoCheleocloeon a new genus of Baetidae (Ephemeroptera)from West Africarsquo Revue drsquoHydrobiologie tropicale 26 213ndash217

Wuillot J and Gillies MT (1994) lsquoDicentroptilum a new genus of Mayflies (BaetidaeEphemeroptera) from Africarsquo Aquatic Insects 16 133ndash140

Zhou CF and Braasch D (2003) lsquoEine neue Gattung und Art der Heptageniidae ausdem ostlichen China (Ephemeroptera)rsquo Entomologische Nachrichten und Berichte 47147ndash151

Zhou CF Gui H and Su CR (2000) lsquoThe first record of the genus Brachycercus in Chinawith description of a new species (Ephemeroptera Caenidae)rsquo Entomologia Sinica 7132ndash134

Zhou CF and Peters JG (2003) lsquoThe nymph of Siphluriscus chinensis and additionalimaginal description A living mayfly with Jurassic origins (Siphluriscidae new familyEphemeroptera)rsquo Florida Entomologist 86 345ndash352

Zhou CF and Zheng LY (2001) lsquoRhithrogena trispina sp n a new species from China(Ephemeroptera Heptageniidae)rsquo Aquatic Insects 23 323ndash326

Zhou CF and Zheng LY (2003) lsquoTwo synonyms and one new species of the genusEphemera from China (Ephemeroptera Ephemeridae)rsquo Acta Zootaxonomica Sinica 28665ndash668

Zhou CF and Zheng LY (2004) lsquoThe genus Prosopistoma from China with description oftwo new species (Ephemeroptera Prosopistomatidae)rsquo Aquatic Insects 26 3ndash8

62 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Lugo-Ortiz CR and McCafferty WP (1999b) lsquoGlobal biodiversity of the mayfly familyBaetidae (Ephemeroptera) a generic perspectiversquo Trends in Entomology 2 45ndash54

Lugo-Ortiz CR and McCafferty WP (1999c) lsquoA new genus of small minnow mayflies(Ephemeroptera Baetidae) with six new species from New Guinea and New BritainrsquoAnnales de Limnologie 35 57ndash70

Lugo-Ortiz CR and McCafferty WP (1999d) lsquoRevision of South American species ofBaetidae (Ephemeroptera) previously placed in Baetis Leach and Pseudocloeon KlapalekrsquoAnnales de Limnologie 35 257ndash262

Lugo-Ortiz CR and McCafferty WP (1999e) lsquoThree new genera of small minnow mayflies(Ephemeroptera Baetidae) from the Andes and Patagoniarsquo Studies on Neotropical Faunaand Environment 34 88ndash104

Lugo-Ortiz CR McCafferty WP and Waltz RD (1999) lsquoDefinition and reorganizationof the genus Pseudocloeon (Ephemeroptera Baetidae) with new species descriptions andcombinationsrsquo Transactions of the American Entomological Society 125 1ndash37

Mayo VK (1973) lsquoFour news species of the genus Baetodes (Ephemeroptera Baetidae)rsquo ThePan-Pacific Entomologist 49 308ndash314

McCafferty WP (1998) lsquoEphemeroptera and the great American interchangersquo Journal of theNorth American Benthological Society 17 1ndash20

McCafferty WP (2000) lsquoNotations on South American Baetidae (Ephemeroptera)rsquoEntomological News 111 375ndash379

McCafferty WP (2009) lsquoMayfly Centralrsquo wwwentmpurdueeduEntomologyresearchmayflymayflyhtml

McCafferty WP and Baumgardner DE (2003) lsquoLugoiops maya a new genus and species ofEphemeroptera (Baetidae) from Central Americarsquo Proceedings of the EntomologicalSociety of Washington 105 397ndash406

McCafferty WP and Sun L (2005) lsquoMystaxiops A new genus of small minnow mayflies(Ephemeroptera Baetidae) from Papua New Guinearsquo Proceedings of the EntomologicalSociety of Washington 107 536ndash542

McDunnough J (1922) lsquoTwo new Canadian May-flies (Ephemeridae)rsquo Canadian Entomol-ogist 53 117ndash120

McDunnough J (1923) lsquoNew Canadian Ephemeridae with notesrsquo Canadian Entomologist55 39ndash50

McDunnough J (1924) lsquoNew Ephemeridae from Illinoisrsquo Canadian Entomologist 56 7ndash9McDunnough J (1925) lsquoNew Canadian Ephemeridae with notes IIIrsquo Canadian Entomol-

ogist 57(168ndash176) 185ndash192McDunnough J (1926) lsquoNew Canadian Ephemeridae with notes IVrsquo Canadian Entomol-

ogist 58 296ndash303McDunnough J (1929) lsquoNotes on North American Ephemeroptera with descriptions of new

species IIrsquo Canadian Entomologist 61 169ndash180McDunnough J (1931) lsquoNew species of North American Ephemeropterarsquo Canadian

Entomologist 63 82ndash93McDunnough J (1932) lsquoNew species of North American Ephemeroptera IIrsquo Canadian

Entomologist 64 209ndash215McDunnough J (1936) lsquoA new Arctic baetid (Ephemeroptera)rsquo Canadian Entomologist 68

32ndash34McDunnough J (1939) lsquoNew British Columbian Ephemeropterarsquo Canadian Entomologist

71 49ndash54Mol AWM (1986) lsquoHarpagobaetis gulosus gen nov sp nov a new mayfly from Suriname

(Ephemeroptera Baetidae)rsquo Zoologische Mededelingen 60 63ndash70Mol AWM (1989) lsquoEchinobaetis phagas gen nov spec nov a new mayfly from Sulawesi

(Ephemeroptera Baetidae)rsquo Zoologische Mededelingen 63 61ndash72Monaghan MT Gattolliat JL Sartori M Elouard JM James H Derleth P Glaizot

O de Moor F and Vogler AP (2005) lsquoTrans-oceanic and endemic origins of the smallminnow mayflies (Ephemeroptera Baetidae) of Madagascarrsquo Proceedings of The RoyalSociety B-Biological Sciences 272 1829ndash1836

Muller-Liebenau I (1980a) lsquoJubabaetis gen n and Platybaetis gen n two new genera of thefamily Baetidae from the Oriental Regionrsquo in Advances in Ephemeroptera Biology edsJF Flannagan and KE Marshall New York Plenum Press pp 103ndash114

58 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Muller-Liebenau I (1980b) lsquoA new species of the genus Platybaetis Muller-Liebenau 1980P bishopi spn from Malaysia (Insecta Ephemeroptera)rsquo Gewasser und Abwasser 666795ndash101

Muller-Liebenau I (1981) lsquoReview of the original material of the baetid genera Baetis andPseudocloeon from the Sunda Islands and the Philippines described by G Ulmer withsome general remarks (Insecta Ephemeroptera)rsquo Mitteilungen der hamburger zoologischerMuseum und Institut 78 197ndash208

Muller-Liebenau I (1982a) lsquoFive new species of Pseudocloeon Klapalek 1905 (FamBaetidae) from the Oriental region (Insecta Ephemeroptera) with some general remarkson Pseudocloeonrsquo Archiv fur Hydrobiologie 95 283ndash298

Muller-Liebenau I (1982b) lsquoA new genus and species of Baetidae from Sri Lanka (Ceylon)Indocloeon primum gen n sp n (Insecta Ephemeroptera)rsquo Aquatic Insects 4 125ndash129

Muller-Liebenau I (1982c) lsquoNew species of the family Baetidae from the Philippines (InsectaEphemeroptera)rsquo Archiv fur Hydrobiologie 94 70ndash82

Muller-Liebenau I (1983) lsquoThree new species of the genus Centroptella Braasch ampSoldan 1980 from Sri Lanka (Insecta Ephemeroptera)rsquo Archiv fur Hydrobiologie 97486ndash500

Muller-Liebenau I (1984a) lsquoBaetidae from Sabah (East Malaysia) (Ephemeroptera)rsquo inProceeding of the fourth international conference on Ephemeroptera eds V LandaT Soldan and M Tonner Bechyne CSAV pp 85ndash99

Muller-Liebenau I (1984b) lsquoNew genera and species of the family Baetidae from West-Malaysia (River Gombak) (Insecta Ephemeroptera)rsquo Spixiana 7 253ndash284

Muller-Liebenau I (1985) lsquoBaetidae from Taiwan with remarks on Baetiella Ueno 1931(Insecta Ephemeroptera)rsquo Archiv fur Hydrobiologie 104 93ndash110

Muller-Liebenau I and Heard WH (1979) lsquoSymbiocloeon a new genus of Baetidaefrom Thailand (Insecta Ephemeroptera)rsquo in Proceedings of the Second InternationalConference on Ephemeroptera eds K Pasternak and R Sowa Warszawa PanstwoweWydawnictwo Naukowe pp 57ndash66

Muller-Liebenau I and Hubbard MD (1985) lsquoBaetidae from Sri Lanka with some generalremarks on the Baetidae of the Oriental region (Insecta Ephemeroptera)rsquo The FloridaEntomologist 68 537ndash561

Muller-Liebenau I and Morihara DK (1982) lsquoIndobaetis A new genus of Baetidae fromSri Lanka (Insecta Ephemeroptera) with two new speciesrsquo Gewasser und Abwasser 686926ndash34

Navas L (1911) lsquoNotas entomologicas Excursiones por los alrededores de Granadarsquo Boletinde la Sociedad Aragonesa de Ciencias Naturales 10 204ndash211

Navas L (1913) lsquoNotas entomologicas Excursiones por los alrededores de Zaragozarsquo Boletinde la Sociedad Aragonesa de Ciencias Naturales 12 75ndash77

Navas L (1917) lsquoNeuropteros nuevos o poco concocidoslsquo Memorias de la Real Academia deCiencias y Artes de Barcelona 13 393ndash395

Navas L (1930) lsquoInsectes du Congo Belge (Serie IV)rsquo Revue de Zoologie et de BotaniqueAfricaines 19 305ndash336

Navas L (1931a) lsquoInsectes du Congo Belge (Serie V)rsquo Revue de Zoologie et de BotaniqueAfricaines 20 257ndash279

Navas L (1931b) lsquoNevropteres et insectes voisins - Chine et pays environnants - 2e SeriersquoNotes drsquoentomologie chinoise 1 1ndash10

Needham JG Traver JR and Hsu Y-S (1935) The biology of mayflies with a systematicaccount of North American species Ithaca NY Comstock Publishing Company

Nieto C (2003a) lsquoEl genero Camelobaetidius (Ephemeroptera Baetidae) en la ArgentinarsquoActa Zoologica Mexicana 88 233ndash255

Nieto C (2003b) lsquoA new species of Guajirolus (Ephemeroptera Baetidae) from Argentinaand description of a new genus from Boliviarsquo in Research Update on Ephemeroptera andPlecoptera ed E Gaino Perugia University of Perugia Italy pp 153ndash158

Nieto C (2004a) lsquoThe genus Baetodes (Ephemeroptera Baetidae) in South America with thedescription of new species from Argentina Bolivia and Perursquo Studies on NeotropicalFauna and Environment 39 63ndash79

Nieto C (2004b) lsquoRedescription of Varipes (Ephemeroptera Baetidae) with the descriptionof new species from Bolivia and Argentinarsquo Aquatic Insects 26 161ndash173

Aquatic Insects 59

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Nieto C (2004c) lsquoSouth American Baetidae (Ephemeroptera) a new generic synonymyrsquoStudies on Neotropical Fauna and Environment 39 95ndash101

Nieto C and Richard B (2008) lsquoThe genus Cloeodes (Ephemeroptera Baetidae) inArgentina with new generic synonymy and new speciesrsquo Zootaxa 1727 1ndash21

Nieto C and Salles FF (2006) lsquoRevision of the genus Paracloeodes (EphemeropteraBaetidae) in South Americarsquo Zootaxa 1ndash33

Nolte U Tietbohl RS and McCafferty WP (1996) lsquoA mayfly from tropical Brazilcapable of tolerating short-term dehydrationrsquo Journal of the North American BenthologicalSociety 15 87ndash94

Park SY Bae YJ and Yoon IB (1996) lsquoRevision of the Baetidae (Ephemeroptera) ofKorea (1) Historical review Acentrella Bengtsson and Baetiella Uenorsquo EntomologicalResearch Bulletin 22 55ndash66

Peters WL (2001) lsquoThe Ephemeroptera of New Caledonia and New Zealandrsquo in Trends inResearch in Ephemeroptera and Plecoptera ed E Domınguez New York KluwerAcademicPlenum Publishers pp 43ndash45

Pictet F-J (1843ndash1845) Histoire naturelle des Insectes Nevropteres Famille des EphemerinesGeneve

Rambur P (1842) Histoire Naturelle des Insectes Nevropteres Paris Librairie Encyclope-dique de Roret

Salles FF (2007) lsquoThe presence of Chane Nieto and Guajirolus Flowers (EphemeropteraBaetidae) in Brazil with the description of a new speciesrsquo Revista Brasileira DeEntomologia 51 404ndash409

Salles FF Andrade MB and Da-Silva ER (2005) lsquoCamelobaetidius francischettii a newspecies of Baetidae (Ephemeroptera) from Brazilrsquo Zootaxa 47ndash53

Salles FF and Batista JD (2004) lsquoThe presence of Varipes Lugo-Ortiz amp McCafferty(Ephemeroptera Baetidae) in Brazil with the description of a new speciesrsquo Zootaxa 4561ndash6

Salles FF Batista JD and Soares Cabette HR (2004) lsquoBaetidae (Insecta Ephemer-optera) de Nova Xavantina Mato Grosso Brasil novos registros e descricao de una novaespecie de Cloeodes Traverrsquo Biota Neotropica 4 1ndash8

Salles FF and Francischetti CN (2004) lsquoCryptonympha dasilvai sp nov (EphemeropteraBaetidae) do Brasilrsquo Neotropical Entomology 33 213ndash216

Salles FF and Polegatto CM (2008) lsquoTwo new species of Baetodes Needham amp Murphy(Ephemeroptera Baetidae) from Brazilrsquo Zootaxa 43ndash50

Sartori M Derleth P and Gattolliat JL (2003) lsquoNew data about the mayflies(Ephemeroptera) from Borneorsquo in Research Update on Ephemeroptera and Plecopteraed E Gaino Perugia University of Perugia Italy pp 403ndash406

Soldan T and Godunko RJ (2006) lsquoBaetis atlanticus n sp a new species of the subgenusRhodobaetis Jacob 2003 from Madeira Portugal (Ephemeroptera Baetidae)rsquo Genus 175ndash17

Soldan T and Thomas AGB (1983a) lsquoBaetis numidicus nsp Ephemeroptere nouveaudrsquoAlgerie (Baetidae)rsquo Annales de Limnologie 19 207ndash211

Soldan T and Thomas AGB (1983b) lsquoNew and little-known species of mayflies(Ephemeroptera) from Algeriarsquo Acta Entomologica Bohemoslovaca 80 356ndash376

Soldan T and Thomas AGB (1985) lsquoCentroptilum dimorphicum sp n a new species ofmayfly (Ephemeroptera Baetidae) from Algeriarsquo Acta Entomologica Bohemoslovaca 82180ndash186

Suter PJ (1979) lsquoA revised key to the Australian genera of mature mayfly(Ephemeroptera) nymphsrsquo Transactions of the Royal Society of South Australia 103 79ndash83

Suter PJ (1986) lsquoThe Ephemeroptera (Mayflies) of South Australiarsquo Records of the SouthAustralian Museum 19 339ndash397

Suter PJ (1997) lsquoPreliminary guide to the identification of nymphs of Australian Baetidmayflies (Insecta Ephemeroptera) found in flowing watersrsquo in Identification Guide No 14Albany Co-Operative Research Center for Freshwater Ecology

Suter PJ (2000) lsquoEdmundsiops hickmani spnov Offadens frater (Tillyard) nov comband description of the nymph of Cloeon tasmaniae Tillyard (EphemeropteraBaetidae) from Tasmaniarsquo Papers and Proceedings of the Royal Society of Tasmania134 63ndash73

60 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Suter PJ (2001) lsquoPlatybaetis gagadjuensis a new species from Northern Australia(Ephemeroptera Baetidae)rsquo in Trends in Research in Ephemeroptera and Plecopteraed E Dominguez New York Kluwer AcademicPlenum Publishers pp 359ndash364

Suter PJ and Pearson MJ (2001) lsquoRedescription of Bungona Harker with new synonymsin the Australian Baetidae (Insecta Ephemeroptera)rsquo Memoirs of the Museum of Victoria58 247ndash254

Thomas A and Gagneur J (1994) lsquoComplements et corrections a la faune desEphemeropteres drsquoAfrique du Nord 6 Alainites sadati n sp drsquoAlgerie (EphemeropteraBaetidae)rsquo Bulletin de la Societe drsquoHistoire Naturelle de Toulouse 130 43ndash45

Thomas A and Peru N (2003) lsquoLes Ephemeres de la Guyane Francaise 6 Description deWaltzoyphius roberti n sp (Ephemeroptera Baetidae)rsquo Bulletin de la Societe drsquoHistoireNaturelle de Toulouse 138 15ndash20 (2002)

Thomas AGB (1986) lsquoEphemeropteres du Sud-Ouest de la France IV Baetis catharus nspdes Pyrenees (Baetidae)rsquo Bulletin de la Societe drsquoHistoire Naturelle de Toulouse 122175ndash179

Thomas AGB (1998) lsquoA provisional checklist of the mayflies of North Africa(Ephemeroptera)rsquo Bulletin de la Societe drsquoHistoire Naturelle de Toulouse 134 13ndash20

Thomas AGB and Dia A (1984) lsquoBaetis baroukianus nsp Ephemeroptere nouveau duLiban (Baetidae)rsquo Bulletin de la Societe drsquoHistoire Naturelle de Toulouse 120 7ndash12

Thomas AGB and Dia A (1985) lsquoBaetis spei nsp Ephemeroptere nouveau du Libandescription et ecologie (Baetidae)rsquo Annales de Limnologie 21 241ndash245

Thomas AGB and Gazagnes G (1984) lsquoBaetis cyrneus n sp Ephemeroptere nouveau deCorse (Baetidae)rsquo Annales de Limnologie 20 199ndash202

Thomas AGB and Sartori M (1989) lsquoMayflies (Insecta Ephemeroptera) of SaudiArabiarsquo Fauna of Saudi Arabia 10 87ndash94

Thomas AGB and Soldan T (1987) lsquoBaetis ingridae n sp Ephemeroptere nouveau deCorse (Baetidae)rsquo Annales de Limnologie 23 23ndash26

Tillyard RJ (1936) lsquoThe trout-food insects of Tasmania Part II- A monograph of themayflies of Tasmaniarsquo Papers and proceedings of the Royal Society of Tasmania 193523ndash59

Tong X and Dudgeon D (2000) lsquoBaetiella (Ephemeroptera Baetidae) in Hong Kong withdescription of a new speciesrsquo Entomological News 111 143ndash148

Tong X and Dudgeon D (2003) lsquoFirst record of the genus Chopralla (EphemeropteraBaetidae) from China and description of a new speciesrsquo The Raffles Bulletin of Zoology51 17ndash19

Traver JR and Edmunds GF (1968) lsquoA revision of the Baetidae with spatulate-clawednymphs (Ephemeroptera)rsquo Pacific Insects 10 629ndash677

Ulmer G (1909) lsquoEphemeriden von Madagaskar und den Comorenrsquo Voeltzkowrsquos Reise inOstrafrika 1903ndash1905 II 365ndash368

Ulmer G (1939) lsquoEintagsfliegen (Ephemeropteren) von den Sunda-Inselnrsquo Archiv furHydrobiologie 16 443ndash692

Walsh BD (1863) lsquoObservations on certain N A Neuroptera by H Hagen M D ofKoenigsberg Prussia translated from the original French ms and published bypermission of the author with notes and descriptions of about twenty new N A speciesof Pseudoneuropterarsquo Proceedings of the Entomological Society of Philadelphia 2167ndash272

Walsh BD (1863) lsquoList of the Pseudoneuroptera of Illinois contained in the cabinet of thewriter with descriptions of over forty new species and notes on their structural affinitiesrsquoProceedings of the Academy of Natural Sciences of Philadelphia 1862 361ndash402

Waltz RD and McCafferty WP (1985) lsquoMoribaetis a new genus of NeotropicalBaetidae (Ephemeroptera)rsquo Proceedings of the Entomological Society of Washington 87239ndash251

Waltz RD and McCafferty WP (1986) lsquoApobaetis etowah (Traver) a new combination inNearctic Baetidae (Ephemeroptera)rsquo Proceedings of the Entomological Society ofWashington 88 191

Waltz RD and McCafferty WP (1987a) lsquoGeneric revision of Cloeodes and description oftwo new genera (Ephemeroptera Baetidae)rsquo Proceedings of the Entomological Society ofWashington 89 177ndash184

Aquatic Insects 61

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Waltz RD and McCafferty WP (1987b) lsquoNew genera of Baetidae for some Nearcticpreviously included in Baetis Leach (Ephemeroptera)rsquo Annals of the Entomological Societyof America 80 667ndash670

Waltz RD and McCafferty WP (1987c) lsquoSystematics of Pseudocloeon AcentrellaBaetiella and Liebebiella new genus (Ephemeroptera Baetidae)rsquo Journal of New YorkEntomological Society 95 553ndash568

Waltz RD and McCafferty WP (1989) lsquoNew species redescriptions and cladisticsof the genus Pseudocentroptiloides (Epherneroptera Baetidae)rsquo Journal of New YorkEntomological Society 97 151ndash158

Waltz RD McCafferty WP and Thomas A (1994) lsquoSystematics of Alainites n genDiphetor Indobaetis Nigrobaetis n stat and Takobia n stat (Ephemeroptera Baetidae)rsquoBulletin de la Societe drsquoHistoire Naturelle de Toulouse 130 33ndash36

Wang T-Q and McCafferty WP (1996) lsquoNew diagnostic characters for the mayfly familyBaetidae (Ephemeroptera)rsquo Entomological News 107 207ndash212

Weyenbergh H (1883) lsquoBijdrage tot de Kennis der zuid-amerikaansche EphemeridenrsquoTijdschrift voor Entomologie 26 159ndash174

Wiersema NA (2000) lsquoA new combination for two North American small minnow mayflies(Ephemeroptera Baetidae)rsquo Entomological News 111 140ndash142

Wiersema NA and McCafferty WP (2004) lsquoNew specific synonyms and records of NorthAmerican Centroptilum and Procloeon (Ephemeroptera Baetidae)rsquo Entomological News115 121ndash128

Wiersema NA Nelson CR and Kuehnl KF (2004) lsquoA new small minnow mayfly(Ephemeroptera Baetidae) from Utah USArsquo Entomological News 115 139ndash145

Wuillot J and Gillies MT (1993) lsquoCheleocloeon a new genus of Baetidae (Ephemeroptera)from West Africarsquo Revue drsquoHydrobiologie tropicale 26 213ndash217

Wuillot J and Gillies MT (1994) lsquoDicentroptilum a new genus of Mayflies (BaetidaeEphemeroptera) from Africarsquo Aquatic Insects 16 133ndash140

Zhou CF and Braasch D (2003) lsquoEine neue Gattung und Art der Heptageniidae ausdem ostlichen China (Ephemeroptera)rsquo Entomologische Nachrichten und Berichte 47147ndash151

Zhou CF Gui H and Su CR (2000) lsquoThe first record of the genus Brachycercus in Chinawith description of a new species (Ephemeroptera Caenidae)rsquo Entomologia Sinica 7132ndash134

Zhou CF and Peters JG (2003) lsquoThe nymph of Siphluriscus chinensis and additionalimaginal description A living mayfly with Jurassic origins (Siphluriscidae new familyEphemeroptera)rsquo Florida Entomologist 86 345ndash352

Zhou CF and Zheng LY (2001) lsquoRhithrogena trispina sp n a new species from China(Ephemeroptera Heptageniidae)rsquo Aquatic Insects 23 323ndash326

Zhou CF and Zheng LY (2003) lsquoTwo synonyms and one new species of the genusEphemera from China (Ephemeroptera Ephemeridae)rsquo Acta Zootaxonomica Sinica 28665ndash668

Zhou CF and Zheng LY (2004) lsquoThe genus Prosopistoma from China with description oftwo new species (Ephemeroptera Prosopistomatidae)rsquo Aquatic Insects 26 3ndash8

62 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Muller-Liebenau I (1980b) lsquoA new species of the genus Platybaetis Muller-Liebenau 1980P bishopi spn from Malaysia (Insecta Ephemeroptera)rsquo Gewasser und Abwasser 666795ndash101

Muller-Liebenau I (1981) lsquoReview of the original material of the baetid genera Baetis andPseudocloeon from the Sunda Islands and the Philippines described by G Ulmer withsome general remarks (Insecta Ephemeroptera)rsquo Mitteilungen der hamburger zoologischerMuseum und Institut 78 197ndash208

Muller-Liebenau I (1982a) lsquoFive new species of Pseudocloeon Klapalek 1905 (FamBaetidae) from the Oriental region (Insecta Ephemeroptera) with some general remarkson Pseudocloeonrsquo Archiv fur Hydrobiologie 95 283ndash298

Muller-Liebenau I (1982b) lsquoA new genus and species of Baetidae from Sri Lanka (Ceylon)Indocloeon primum gen n sp n (Insecta Ephemeroptera)rsquo Aquatic Insects 4 125ndash129

Muller-Liebenau I (1982c) lsquoNew species of the family Baetidae from the Philippines (InsectaEphemeroptera)rsquo Archiv fur Hydrobiologie 94 70ndash82

Muller-Liebenau I (1983) lsquoThree new species of the genus Centroptella Braasch ampSoldan 1980 from Sri Lanka (Insecta Ephemeroptera)rsquo Archiv fur Hydrobiologie 97486ndash500

Muller-Liebenau I (1984a) lsquoBaetidae from Sabah (East Malaysia) (Ephemeroptera)rsquo inProceeding of the fourth international conference on Ephemeroptera eds V LandaT Soldan and M Tonner Bechyne CSAV pp 85ndash99

Muller-Liebenau I (1984b) lsquoNew genera and species of the family Baetidae from West-Malaysia (River Gombak) (Insecta Ephemeroptera)rsquo Spixiana 7 253ndash284

Muller-Liebenau I (1985) lsquoBaetidae from Taiwan with remarks on Baetiella Ueno 1931(Insecta Ephemeroptera)rsquo Archiv fur Hydrobiologie 104 93ndash110

Muller-Liebenau I and Heard WH (1979) lsquoSymbiocloeon a new genus of Baetidaefrom Thailand (Insecta Ephemeroptera)rsquo in Proceedings of the Second InternationalConference on Ephemeroptera eds K Pasternak and R Sowa Warszawa PanstwoweWydawnictwo Naukowe pp 57ndash66

Muller-Liebenau I and Hubbard MD (1985) lsquoBaetidae from Sri Lanka with some generalremarks on the Baetidae of the Oriental region (Insecta Ephemeroptera)rsquo The FloridaEntomologist 68 537ndash561

Muller-Liebenau I and Morihara DK (1982) lsquoIndobaetis A new genus of Baetidae fromSri Lanka (Insecta Ephemeroptera) with two new speciesrsquo Gewasser und Abwasser 686926ndash34

Navas L (1911) lsquoNotas entomologicas Excursiones por los alrededores de Granadarsquo Boletinde la Sociedad Aragonesa de Ciencias Naturales 10 204ndash211

Navas L (1913) lsquoNotas entomologicas Excursiones por los alrededores de Zaragozarsquo Boletinde la Sociedad Aragonesa de Ciencias Naturales 12 75ndash77

Navas L (1917) lsquoNeuropteros nuevos o poco concocidoslsquo Memorias de la Real Academia deCiencias y Artes de Barcelona 13 393ndash395

Navas L (1930) lsquoInsectes du Congo Belge (Serie IV)rsquo Revue de Zoologie et de BotaniqueAfricaines 19 305ndash336

Navas L (1931a) lsquoInsectes du Congo Belge (Serie V)rsquo Revue de Zoologie et de BotaniqueAfricaines 20 257ndash279

Navas L (1931b) lsquoNevropteres et insectes voisins - Chine et pays environnants - 2e SeriersquoNotes drsquoentomologie chinoise 1 1ndash10

Needham JG Traver JR and Hsu Y-S (1935) The biology of mayflies with a systematicaccount of North American species Ithaca NY Comstock Publishing Company

Nieto C (2003a) lsquoEl genero Camelobaetidius (Ephemeroptera Baetidae) en la ArgentinarsquoActa Zoologica Mexicana 88 233ndash255

Nieto C (2003b) lsquoA new species of Guajirolus (Ephemeroptera Baetidae) from Argentinaand description of a new genus from Boliviarsquo in Research Update on Ephemeroptera andPlecoptera ed E Gaino Perugia University of Perugia Italy pp 153ndash158

Nieto C (2004a) lsquoThe genus Baetodes (Ephemeroptera Baetidae) in South America with thedescription of new species from Argentina Bolivia and Perursquo Studies on NeotropicalFauna and Environment 39 63ndash79

Nieto C (2004b) lsquoRedescription of Varipes (Ephemeroptera Baetidae) with the descriptionof new species from Bolivia and Argentinarsquo Aquatic Insects 26 161ndash173

Aquatic Insects 59

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Nieto C (2004c) lsquoSouth American Baetidae (Ephemeroptera) a new generic synonymyrsquoStudies on Neotropical Fauna and Environment 39 95ndash101

Nieto C and Richard B (2008) lsquoThe genus Cloeodes (Ephemeroptera Baetidae) inArgentina with new generic synonymy and new speciesrsquo Zootaxa 1727 1ndash21

Nieto C and Salles FF (2006) lsquoRevision of the genus Paracloeodes (EphemeropteraBaetidae) in South Americarsquo Zootaxa 1ndash33

Nolte U Tietbohl RS and McCafferty WP (1996) lsquoA mayfly from tropical Brazilcapable of tolerating short-term dehydrationrsquo Journal of the North American BenthologicalSociety 15 87ndash94

Park SY Bae YJ and Yoon IB (1996) lsquoRevision of the Baetidae (Ephemeroptera) ofKorea (1) Historical review Acentrella Bengtsson and Baetiella Uenorsquo EntomologicalResearch Bulletin 22 55ndash66

Peters WL (2001) lsquoThe Ephemeroptera of New Caledonia and New Zealandrsquo in Trends inResearch in Ephemeroptera and Plecoptera ed E Domınguez New York KluwerAcademicPlenum Publishers pp 43ndash45

Pictet F-J (1843ndash1845) Histoire naturelle des Insectes Nevropteres Famille des EphemerinesGeneve

Rambur P (1842) Histoire Naturelle des Insectes Nevropteres Paris Librairie Encyclope-dique de Roret

Salles FF (2007) lsquoThe presence of Chane Nieto and Guajirolus Flowers (EphemeropteraBaetidae) in Brazil with the description of a new speciesrsquo Revista Brasileira DeEntomologia 51 404ndash409

Salles FF Andrade MB and Da-Silva ER (2005) lsquoCamelobaetidius francischettii a newspecies of Baetidae (Ephemeroptera) from Brazilrsquo Zootaxa 47ndash53

Salles FF and Batista JD (2004) lsquoThe presence of Varipes Lugo-Ortiz amp McCafferty(Ephemeroptera Baetidae) in Brazil with the description of a new speciesrsquo Zootaxa 4561ndash6

Salles FF Batista JD and Soares Cabette HR (2004) lsquoBaetidae (Insecta Ephemer-optera) de Nova Xavantina Mato Grosso Brasil novos registros e descricao de una novaespecie de Cloeodes Traverrsquo Biota Neotropica 4 1ndash8

Salles FF and Francischetti CN (2004) lsquoCryptonympha dasilvai sp nov (EphemeropteraBaetidae) do Brasilrsquo Neotropical Entomology 33 213ndash216

Salles FF and Polegatto CM (2008) lsquoTwo new species of Baetodes Needham amp Murphy(Ephemeroptera Baetidae) from Brazilrsquo Zootaxa 43ndash50

Sartori M Derleth P and Gattolliat JL (2003) lsquoNew data about the mayflies(Ephemeroptera) from Borneorsquo in Research Update on Ephemeroptera and Plecopteraed E Gaino Perugia University of Perugia Italy pp 403ndash406

Soldan T and Godunko RJ (2006) lsquoBaetis atlanticus n sp a new species of the subgenusRhodobaetis Jacob 2003 from Madeira Portugal (Ephemeroptera Baetidae)rsquo Genus 175ndash17

Soldan T and Thomas AGB (1983a) lsquoBaetis numidicus nsp Ephemeroptere nouveaudrsquoAlgerie (Baetidae)rsquo Annales de Limnologie 19 207ndash211

Soldan T and Thomas AGB (1983b) lsquoNew and little-known species of mayflies(Ephemeroptera) from Algeriarsquo Acta Entomologica Bohemoslovaca 80 356ndash376

Soldan T and Thomas AGB (1985) lsquoCentroptilum dimorphicum sp n a new species ofmayfly (Ephemeroptera Baetidae) from Algeriarsquo Acta Entomologica Bohemoslovaca 82180ndash186

Suter PJ (1979) lsquoA revised key to the Australian genera of mature mayfly(Ephemeroptera) nymphsrsquo Transactions of the Royal Society of South Australia 103 79ndash83

Suter PJ (1986) lsquoThe Ephemeroptera (Mayflies) of South Australiarsquo Records of the SouthAustralian Museum 19 339ndash397

Suter PJ (1997) lsquoPreliminary guide to the identification of nymphs of Australian Baetidmayflies (Insecta Ephemeroptera) found in flowing watersrsquo in Identification Guide No 14Albany Co-Operative Research Center for Freshwater Ecology

Suter PJ (2000) lsquoEdmundsiops hickmani spnov Offadens frater (Tillyard) nov comband description of the nymph of Cloeon tasmaniae Tillyard (EphemeropteraBaetidae) from Tasmaniarsquo Papers and Proceedings of the Royal Society of Tasmania134 63ndash73

60 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Suter PJ (2001) lsquoPlatybaetis gagadjuensis a new species from Northern Australia(Ephemeroptera Baetidae)rsquo in Trends in Research in Ephemeroptera and Plecopteraed E Dominguez New York Kluwer AcademicPlenum Publishers pp 359ndash364

Suter PJ and Pearson MJ (2001) lsquoRedescription of Bungona Harker with new synonymsin the Australian Baetidae (Insecta Ephemeroptera)rsquo Memoirs of the Museum of Victoria58 247ndash254

Thomas A and Gagneur J (1994) lsquoComplements et corrections a la faune desEphemeropteres drsquoAfrique du Nord 6 Alainites sadati n sp drsquoAlgerie (EphemeropteraBaetidae)rsquo Bulletin de la Societe drsquoHistoire Naturelle de Toulouse 130 43ndash45

Thomas A and Peru N (2003) lsquoLes Ephemeres de la Guyane Francaise 6 Description deWaltzoyphius roberti n sp (Ephemeroptera Baetidae)rsquo Bulletin de la Societe drsquoHistoireNaturelle de Toulouse 138 15ndash20 (2002)

Thomas AGB (1986) lsquoEphemeropteres du Sud-Ouest de la France IV Baetis catharus nspdes Pyrenees (Baetidae)rsquo Bulletin de la Societe drsquoHistoire Naturelle de Toulouse 122175ndash179

Thomas AGB (1998) lsquoA provisional checklist of the mayflies of North Africa(Ephemeroptera)rsquo Bulletin de la Societe drsquoHistoire Naturelle de Toulouse 134 13ndash20

Thomas AGB and Dia A (1984) lsquoBaetis baroukianus nsp Ephemeroptere nouveau duLiban (Baetidae)rsquo Bulletin de la Societe drsquoHistoire Naturelle de Toulouse 120 7ndash12

Thomas AGB and Dia A (1985) lsquoBaetis spei nsp Ephemeroptere nouveau du Libandescription et ecologie (Baetidae)rsquo Annales de Limnologie 21 241ndash245

Thomas AGB and Gazagnes G (1984) lsquoBaetis cyrneus n sp Ephemeroptere nouveau deCorse (Baetidae)rsquo Annales de Limnologie 20 199ndash202

Thomas AGB and Sartori M (1989) lsquoMayflies (Insecta Ephemeroptera) of SaudiArabiarsquo Fauna of Saudi Arabia 10 87ndash94

Thomas AGB and Soldan T (1987) lsquoBaetis ingridae n sp Ephemeroptere nouveau deCorse (Baetidae)rsquo Annales de Limnologie 23 23ndash26

Tillyard RJ (1936) lsquoThe trout-food insects of Tasmania Part II- A monograph of themayflies of Tasmaniarsquo Papers and proceedings of the Royal Society of Tasmania 193523ndash59

Tong X and Dudgeon D (2000) lsquoBaetiella (Ephemeroptera Baetidae) in Hong Kong withdescription of a new speciesrsquo Entomological News 111 143ndash148

Tong X and Dudgeon D (2003) lsquoFirst record of the genus Chopralla (EphemeropteraBaetidae) from China and description of a new speciesrsquo The Raffles Bulletin of Zoology51 17ndash19

Traver JR and Edmunds GF (1968) lsquoA revision of the Baetidae with spatulate-clawednymphs (Ephemeroptera)rsquo Pacific Insects 10 629ndash677

Ulmer G (1909) lsquoEphemeriden von Madagaskar und den Comorenrsquo Voeltzkowrsquos Reise inOstrafrika 1903ndash1905 II 365ndash368

Ulmer G (1939) lsquoEintagsfliegen (Ephemeropteren) von den Sunda-Inselnrsquo Archiv furHydrobiologie 16 443ndash692

Walsh BD (1863) lsquoObservations on certain N A Neuroptera by H Hagen M D ofKoenigsberg Prussia translated from the original French ms and published bypermission of the author with notes and descriptions of about twenty new N A speciesof Pseudoneuropterarsquo Proceedings of the Entomological Society of Philadelphia 2167ndash272

Walsh BD (1863) lsquoList of the Pseudoneuroptera of Illinois contained in the cabinet of thewriter with descriptions of over forty new species and notes on their structural affinitiesrsquoProceedings of the Academy of Natural Sciences of Philadelphia 1862 361ndash402

Waltz RD and McCafferty WP (1985) lsquoMoribaetis a new genus of NeotropicalBaetidae (Ephemeroptera)rsquo Proceedings of the Entomological Society of Washington 87239ndash251

Waltz RD and McCafferty WP (1986) lsquoApobaetis etowah (Traver) a new combination inNearctic Baetidae (Ephemeroptera)rsquo Proceedings of the Entomological Society ofWashington 88 191

Waltz RD and McCafferty WP (1987a) lsquoGeneric revision of Cloeodes and description oftwo new genera (Ephemeroptera Baetidae)rsquo Proceedings of the Entomological Society ofWashington 89 177ndash184

Aquatic Insects 61

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Waltz RD and McCafferty WP (1987b) lsquoNew genera of Baetidae for some Nearcticpreviously included in Baetis Leach (Ephemeroptera)rsquo Annals of the Entomological Societyof America 80 667ndash670

Waltz RD and McCafferty WP (1987c) lsquoSystematics of Pseudocloeon AcentrellaBaetiella and Liebebiella new genus (Ephemeroptera Baetidae)rsquo Journal of New YorkEntomological Society 95 553ndash568

Waltz RD and McCafferty WP (1989) lsquoNew species redescriptions and cladisticsof the genus Pseudocentroptiloides (Epherneroptera Baetidae)rsquo Journal of New YorkEntomological Society 97 151ndash158

Waltz RD McCafferty WP and Thomas A (1994) lsquoSystematics of Alainites n genDiphetor Indobaetis Nigrobaetis n stat and Takobia n stat (Ephemeroptera Baetidae)rsquoBulletin de la Societe drsquoHistoire Naturelle de Toulouse 130 33ndash36

Wang T-Q and McCafferty WP (1996) lsquoNew diagnostic characters for the mayfly familyBaetidae (Ephemeroptera)rsquo Entomological News 107 207ndash212

Weyenbergh H (1883) lsquoBijdrage tot de Kennis der zuid-amerikaansche EphemeridenrsquoTijdschrift voor Entomologie 26 159ndash174

Wiersema NA (2000) lsquoA new combination for two North American small minnow mayflies(Ephemeroptera Baetidae)rsquo Entomological News 111 140ndash142

Wiersema NA and McCafferty WP (2004) lsquoNew specific synonyms and records of NorthAmerican Centroptilum and Procloeon (Ephemeroptera Baetidae)rsquo Entomological News115 121ndash128

Wiersema NA Nelson CR and Kuehnl KF (2004) lsquoA new small minnow mayfly(Ephemeroptera Baetidae) from Utah USArsquo Entomological News 115 139ndash145

Wuillot J and Gillies MT (1993) lsquoCheleocloeon a new genus of Baetidae (Ephemeroptera)from West Africarsquo Revue drsquoHydrobiologie tropicale 26 213ndash217

Wuillot J and Gillies MT (1994) lsquoDicentroptilum a new genus of Mayflies (BaetidaeEphemeroptera) from Africarsquo Aquatic Insects 16 133ndash140

Zhou CF and Braasch D (2003) lsquoEine neue Gattung und Art der Heptageniidae ausdem ostlichen China (Ephemeroptera)rsquo Entomologische Nachrichten und Berichte 47147ndash151

Zhou CF Gui H and Su CR (2000) lsquoThe first record of the genus Brachycercus in Chinawith description of a new species (Ephemeroptera Caenidae)rsquo Entomologia Sinica 7132ndash134

Zhou CF and Peters JG (2003) lsquoThe nymph of Siphluriscus chinensis and additionalimaginal description A living mayfly with Jurassic origins (Siphluriscidae new familyEphemeroptera)rsquo Florida Entomologist 86 345ndash352

Zhou CF and Zheng LY (2001) lsquoRhithrogena trispina sp n a new species from China(Ephemeroptera Heptageniidae)rsquo Aquatic Insects 23 323ndash326

Zhou CF and Zheng LY (2003) lsquoTwo synonyms and one new species of the genusEphemera from China (Ephemeroptera Ephemeridae)rsquo Acta Zootaxonomica Sinica 28665ndash668

Zhou CF and Zheng LY (2004) lsquoThe genus Prosopistoma from China with description oftwo new species (Ephemeroptera Prosopistomatidae)rsquo Aquatic Insects 26 3ndash8

62 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Nieto C (2004c) lsquoSouth American Baetidae (Ephemeroptera) a new generic synonymyrsquoStudies on Neotropical Fauna and Environment 39 95ndash101

Nieto C and Richard B (2008) lsquoThe genus Cloeodes (Ephemeroptera Baetidae) inArgentina with new generic synonymy and new speciesrsquo Zootaxa 1727 1ndash21

Nieto C and Salles FF (2006) lsquoRevision of the genus Paracloeodes (EphemeropteraBaetidae) in South Americarsquo Zootaxa 1ndash33

Nolte U Tietbohl RS and McCafferty WP (1996) lsquoA mayfly from tropical Brazilcapable of tolerating short-term dehydrationrsquo Journal of the North American BenthologicalSociety 15 87ndash94

Park SY Bae YJ and Yoon IB (1996) lsquoRevision of the Baetidae (Ephemeroptera) ofKorea (1) Historical review Acentrella Bengtsson and Baetiella Uenorsquo EntomologicalResearch Bulletin 22 55ndash66

Peters WL (2001) lsquoThe Ephemeroptera of New Caledonia and New Zealandrsquo in Trends inResearch in Ephemeroptera and Plecoptera ed E Domınguez New York KluwerAcademicPlenum Publishers pp 43ndash45

Pictet F-J (1843ndash1845) Histoire naturelle des Insectes Nevropteres Famille des EphemerinesGeneve

Rambur P (1842) Histoire Naturelle des Insectes Nevropteres Paris Librairie Encyclope-dique de Roret

Salles FF (2007) lsquoThe presence of Chane Nieto and Guajirolus Flowers (EphemeropteraBaetidae) in Brazil with the description of a new speciesrsquo Revista Brasileira DeEntomologia 51 404ndash409

Salles FF Andrade MB and Da-Silva ER (2005) lsquoCamelobaetidius francischettii a newspecies of Baetidae (Ephemeroptera) from Brazilrsquo Zootaxa 47ndash53

Salles FF and Batista JD (2004) lsquoThe presence of Varipes Lugo-Ortiz amp McCafferty(Ephemeroptera Baetidae) in Brazil with the description of a new speciesrsquo Zootaxa 4561ndash6

Salles FF Batista JD and Soares Cabette HR (2004) lsquoBaetidae (Insecta Ephemer-optera) de Nova Xavantina Mato Grosso Brasil novos registros e descricao de una novaespecie de Cloeodes Traverrsquo Biota Neotropica 4 1ndash8

Salles FF and Francischetti CN (2004) lsquoCryptonympha dasilvai sp nov (EphemeropteraBaetidae) do Brasilrsquo Neotropical Entomology 33 213ndash216

Salles FF and Polegatto CM (2008) lsquoTwo new species of Baetodes Needham amp Murphy(Ephemeroptera Baetidae) from Brazilrsquo Zootaxa 43ndash50

Sartori M Derleth P and Gattolliat JL (2003) lsquoNew data about the mayflies(Ephemeroptera) from Borneorsquo in Research Update on Ephemeroptera and Plecopteraed E Gaino Perugia University of Perugia Italy pp 403ndash406

Soldan T and Godunko RJ (2006) lsquoBaetis atlanticus n sp a new species of the subgenusRhodobaetis Jacob 2003 from Madeira Portugal (Ephemeroptera Baetidae)rsquo Genus 175ndash17

Soldan T and Thomas AGB (1983a) lsquoBaetis numidicus nsp Ephemeroptere nouveaudrsquoAlgerie (Baetidae)rsquo Annales de Limnologie 19 207ndash211

Soldan T and Thomas AGB (1983b) lsquoNew and little-known species of mayflies(Ephemeroptera) from Algeriarsquo Acta Entomologica Bohemoslovaca 80 356ndash376

Soldan T and Thomas AGB (1985) lsquoCentroptilum dimorphicum sp n a new species ofmayfly (Ephemeroptera Baetidae) from Algeriarsquo Acta Entomologica Bohemoslovaca 82180ndash186

Suter PJ (1979) lsquoA revised key to the Australian genera of mature mayfly(Ephemeroptera) nymphsrsquo Transactions of the Royal Society of South Australia 103 79ndash83

Suter PJ (1986) lsquoThe Ephemeroptera (Mayflies) of South Australiarsquo Records of the SouthAustralian Museum 19 339ndash397

Suter PJ (1997) lsquoPreliminary guide to the identification of nymphs of Australian Baetidmayflies (Insecta Ephemeroptera) found in flowing watersrsquo in Identification Guide No 14Albany Co-Operative Research Center for Freshwater Ecology

Suter PJ (2000) lsquoEdmundsiops hickmani spnov Offadens frater (Tillyard) nov comband description of the nymph of Cloeon tasmaniae Tillyard (EphemeropteraBaetidae) from Tasmaniarsquo Papers and Proceedings of the Royal Society of Tasmania134 63ndash73

60 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Suter PJ (2001) lsquoPlatybaetis gagadjuensis a new species from Northern Australia(Ephemeroptera Baetidae)rsquo in Trends in Research in Ephemeroptera and Plecopteraed E Dominguez New York Kluwer AcademicPlenum Publishers pp 359ndash364

Suter PJ and Pearson MJ (2001) lsquoRedescription of Bungona Harker with new synonymsin the Australian Baetidae (Insecta Ephemeroptera)rsquo Memoirs of the Museum of Victoria58 247ndash254

Thomas A and Gagneur J (1994) lsquoComplements et corrections a la faune desEphemeropteres drsquoAfrique du Nord 6 Alainites sadati n sp drsquoAlgerie (EphemeropteraBaetidae)rsquo Bulletin de la Societe drsquoHistoire Naturelle de Toulouse 130 43ndash45

Thomas A and Peru N (2003) lsquoLes Ephemeres de la Guyane Francaise 6 Description deWaltzoyphius roberti n sp (Ephemeroptera Baetidae)rsquo Bulletin de la Societe drsquoHistoireNaturelle de Toulouse 138 15ndash20 (2002)

Thomas AGB (1986) lsquoEphemeropteres du Sud-Ouest de la France IV Baetis catharus nspdes Pyrenees (Baetidae)rsquo Bulletin de la Societe drsquoHistoire Naturelle de Toulouse 122175ndash179

Thomas AGB (1998) lsquoA provisional checklist of the mayflies of North Africa(Ephemeroptera)rsquo Bulletin de la Societe drsquoHistoire Naturelle de Toulouse 134 13ndash20

Thomas AGB and Dia A (1984) lsquoBaetis baroukianus nsp Ephemeroptere nouveau duLiban (Baetidae)rsquo Bulletin de la Societe drsquoHistoire Naturelle de Toulouse 120 7ndash12

Thomas AGB and Dia A (1985) lsquoBaetis spei nsp Ephemeroptere nouveau du Libandescription et ecologie (Baetidae)rsquo Annales de Limnologie 21 241ndash245

Thomas AGB and Gazagnes G (1984) lsquoBaetis cyrneus n sp Ephemeroptere nouveau deCorse (Baetidae)rsquo Annales de Limnologie 20 199ndash202

Thomas AGB and Sartori M (1989) lsquoMayflies (Insecta Ephemeroptera) of SaudiArabiarsquo Fauna of Saudi Arabia 10 87ndash94

Thomas AGB and Soldan T (1987) lsquoBaetis ingridae n sp Ephemeroptere nouveau deCorse (Baetidae)rsquo Annales de Limnologie 23 23ndash26

Tillyard RJ (1936) lsquoThe trout-food insects of Tasmania Part II- A monograph of themayflies of Tasmaniarsquo Papers and proceedings of the Royal Society of Tasmania 193523ndash59

Tong X and Dudgeon D (2000) lsquoBaetiella (Ephemeroptera Baetidae) in Hong Kong withdescription of a new speciesrsquo Entomological News 111 143ndash148

Tong X and Dudgeon D (2003) lsquoFirst record of the genus Chopralla (EphemeropteraBaetidae) from China and description of a new speciesrsquo The Raffles Bulletin of Zoology51 17ndash19

Traver JR and Edmunds GF (1968) lsquoA revision of the Baetidae with spatulate-clawednymphs (Ephemeroptera)rsquo Pacific Insects 10 629ndash677

Ulmer G (1909) lsquoEphemeriden von Madagaskar und den Comorenrsquo Voeltzkowrsquos Reise inOstrafrika 1903ndash1905 II 365ndash368

Ulmer G (1939) lsquoEintagsfliegen (Ephemeropteren) von den Sunda-Inselnrsquo Archiv furHydrobiologie 16 443ndash692

Walsh BD (1863) lsquoObservations on certain N A Neuroptera by H Hagen M D ofKoenigsberg Prussia translated from the original French ms and published bypermission of the author with notes and descriptions of about twenty new N A speciesof Pseudoneuropterarsquo Proceedings of the Entomological Society of Philadelphia 2167ndash272

Walsh BD (1863) lsquoList of the Pseudoneuroptera of Illinois contained in the cabinet of thewriter with descriptions of over forty new species and notes on their structural affinitiesrsquoProceedings of the Academy of Natural Sciences of Philadelphia 1862 361ndash402

Waltz RD and McCafferty WP (1985) lsquoMoribaetis a new genus of NeotropicalBaetidae (Ephemeroptera)rsquo Proceedings of the Entomological Society of Washington 87239ndash251

Waltz RD and McCafferty WP (1986) lsquoApobaetis etowah (Traver) a new combination inNearctic Baetidae (Ephemeroptera)rsquo Proceedings of the Entomological Society ofWashington 88 191

Waltz RD and McCafferty WP (1987a) lsquoGeneric revision of Cloeodes and description oftwo new genera (Ephemeroptera Baetidae)rsquo Proceedings of the Entomological Society ofWashington 89 177ndash184

Aquatic Insects 61

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Waltz RD and McCafferty WP (1987b) lsquoNew genera of Baetidae for some Nearcticpreviously included in Baetis Leach (Ephemeroptera)rsquo Annals of the Entomological Societyof America 80 667ndash670

Waltz RD and McCafferty WP (1987c) lsquoSystematics of Pseudocloeon AcentrellaBaetiella and Liebebiella new genus (Ephemeroptera Baetidae)rsquo Journal of New YorkEntomological Society 95 553ndash568

Waltz RD and McCafferty WP (1989) lsquoNew species redescriptions and cladisticsof the genus Pseudocentroptiloides (Epherneroptera Baetidae)rsquo Journal of New YorkEntomological Society 97 151ndash158

Waltz RD McCafferty WP and Thomas A (1994) lsquoSystematics of Alainites n genDiphetor Indobaetis Nigrobaetis n stat and Takobia n stat (Ephemeroptera Baetidae)rsquoBulletin de la Societe drsquoHistoire Naturelle de Toulouse 130 33ndash36

Wang T-Q and McCafferty WP (1996) lsquoNew diagnostic characters for the mayfly familyBaetidae (Ephemeroptera)rsquo Entomological News 107 207ndash212

Weyenbergh H (1883) lsquoBijdrage tot de Kennis der zuid-amerikaansche EphemeridenrsquoTijdschrift voor Entomologie 26 159ndash174

Wiersema NA (2000) lsquoA new combination for two North American small minnow mayflies(Ephemeroptera Baetidae)rsquo Entomological News 111 140ndash142

Wiersema NA and McCafferty WP (2004) lsquoNew specific synonyms and records of NorthAmerican Centroptilum and Procloeon (Ephemeroptera Baetidae)rsquo Entomological News115 121ndash128

Wiersema NA Nelson CR and Kuehnl KF (2004) lsquoA new small minnow mayfly(Ephemeroptera Baetidae) from Utah USArsquo Entomological News 115 139ndash145

Wuillot J and Gillies MT (1993) lsquoCheleocloeon a new genus of Baetidae (Ephemeroptera)from West Africarsquo Revue drsquoHydrobiologie tropicale 26 213ndash217

Wuillot J and Gillies MT (1994) lsquoDicentroptilum a new genus of Mayflies (BaetidaeEphemeroptera) from Africarsquo Aquatic Insects 16 133ndash140

Zhou CF and Braasch D (2003) lsquoEine neue Gattung und Art der Heptageniidae ausdem ostlichen China (Ephemeroptera)rsquo Entomologische Nachrichten und Berichte 47147ndash151

Zhou CF Gui H and Su CR (2000) lsquoThe first record of the genus Brachycercus in Chinawith description of a new species (Ephemeroptera Caenidae)rsquo Entomologia Sinica 7132ndash134

Zhou CF and Peters JG (2003) lsquoThe nymph of Siphluriscus chinensis and additionalimaginal description A living mayfly with Jurassic origins (Siphluriscidae new familyEphemeroptera)rsquo Florida Entomologist 86 345ndash352

Zhou CF and Zheng LY (2001) lsquoRhithrogena trispina sp n a new species from China(Ephemeroptera Heptageniidae)rsquo Aquatic Insects 23 323ndash326

Zhou CF and Zheng LY (2003) lsquoTwo synonyms and one new species of the genusEphemera from China (Ephemeroptera Ephemeridae)rsquo Acta Zootaxonomica Sinica 28665ndash668

Zhou CF and Zheng LY (2004) lsquoThe genus Prosopistoma from China with description oftwo new species (Ephemeroptera Prosopistomatidae)rsquo Aquatic Insects 26 3ndash8

62 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Suter PJ (2001) lsquoPlatybaetis gagadjuensis a new species from Northern Australia(Ephemeroptera Baetidae)rsquo in Trends in Research in Ephemeroptera and Plecopteraed E Dominguez New York Kluwer AcademicPlenum Publishers pp 359ndash364

Suter PJ and Pearson MJ (2001) lsquoRedescription of Bungona Harker with new synonymsin the Australian Baetidae (Insecta Ephemeroptera)rsquo Memoirs of the Museum of Victoria58 247ndash254

Thomas A and Gagneur J (1994) lsquoComplements et corrections a la faune desEphemeropteres drsquoAfrique du Nord 6 Alainites sadati n sp drsquoAlgerie (EphemeropteraBaetidae)rsquo Bulletin de la Societe drsquoHistoire Naturelle de Toulouse 130 43ndash45

Thomas A and Peru N (2003) lsquoLes Ephemeres de la Guyane Francaise 6 Description deWaltzoyphius roberti n sp (Ephemeroptera Baetidae)rsquo Bulletin de la Societe drsquoHistoireNaturelle de Toulouse 138 15ndash20 (2002)

Thomas AGB (1986) lsquoEphemeropteres du Sud-Ouest de la France IV Baetis catharus nspdes Pyrenees (Baetidae)rsquo Bulletin de la Societe drsquoHistoire Naturelle de Toulouse 122175ndash179

Thomas AGB (1998) lsquoA provisional checklist of the mayflies of North Africa(Ephemeroptera)rsquo Bulletin de la Societe drsquoHistoire Naturelle de Toulouse 134 13ndash20

Thomas AGB and Dia A (1984) lsquoBaetis baroukianus nsp Ephemeroptere nouveau duLiban (Baetidae)rsquo Bulletin de la Societe drsquoHistoire Naturelle de Toulouse 120 7ndash12

Thomas AGB and Dia A (1985) lsquoBaetis spei nsp Ephemeroptere nouveau du Libandescription et ecologie (Baetidae)rsquo Annales de Limnologie 21 241ndash245

Thomas AGB and Gazagnes G (1984) lsquoBaetis cyrneus n sp Ephemeroptere nouveau deCorse (Baetidae)rsquo Annales de Limnologie 20 199ndash202

Thomas AGB and Sartori M (1989) lsquoMayflies (Insecta Ephemeroptera) of SaudiArabiarsquo Fauna of Saudi Arabia 10 87ndash94

Thomas AGB and Soldan T (1987) lsquoBaetis ingridae n sp Ephemeroptere nouveau deCorse (Baetidae)rsquo Annales de Limnologie 23 23ndash26

Tillyard RJ (1936) lsquoThe trout-food insects of Tasmania Part II- A monograph of themayflies of Tasmaniarsquo Papers and proceedings of the Royal Society of Tasmania 193523ndash59

Tong X and Dudgeon D (2000) lsquoBaetiella (Ephemeroptera Baetidae) in Hong Kong withdescription of a new speciesrsquo Entomological News 111 143ndash148

Tong X and Dudgeon D (2003) lsquoFirst record of the genus Chopralla (EphemeropteraBaetidae) from China and description of a new speciesrsquo The Raffles Bulletin of Zoology51 17ndash19

Traver JR and Edmunds GF (1968) lsquoA revision of the Baetidae with spatulate-clawednymphs (Ephemeroptera)rsquo Pacific Insects 10 629ndash677

Ulmer G (1909) lsquoEphemeriden von Madagaskar und den Comorenrsquo Voeltzkowrsquos Reise inOstrafrika 1903ndash1905 II 365ndash368

Ulmer G (1939) lsquoEintagsfliegen (Ephemeropteren) von den Sunda-Inselnrsquo Archiv furHydrobiologie 16 443ndash692

Walsh BD (1863) lsquoObservations on certain N A Neuroptera by H Hagen M D ofKoenigsberg Prussia translated from the original French ms and published bypermission of the author with notes and descriptions of about twenty new N A speciesof Pseudoneuropterarsquo Proceedings of the Entomological Society of Philadelphia 2167ndash272

Walsh BD (1863) lsquoList of the Pseudoneuroptera of Illinois contained in the cabinet of thewriter with descriptions of over forty new species and notes on their structural affinitiesrsquoProceedings of the Academy of Natural Sciences of Philadelphia 1862 361ndash402

Waltz RD and McCafferty WP (1985) lsquoMoribaetis a new genus of NeotropicalBaetidae (Ephemeroptera)rsquo Proceedings of the Entomological Society of Washington 87239ndash251

Waltz RD and McCafferty WP (1986) lsquoApobaetis etowah (Traver) a new combination inNearctic Baetidae (Ephemeroptera)rsquo Proceedings of the Entomological Society ofWashington 88 191

Waltz RD and McCafferty WP (1987a) lsquoGeneric revision of Cloeodes and description oftwo new genera (Ephemeroptera Baetidae)rsquo Proceedings of the Entomological Society ofWashington 89 177ndash184

Aquatic Insects 61

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Waltz RD and McCafferty WP (1987b) lsquoNew genera of Baetidae for some Nearcticpreviously included in Baetis Leach (Ephemeroptera)rsquo Annals of the Entomological Societyof America 80 667ndash670

Waltz RD and McCafferty WP (1987c) lsquoSystematics of Pseudocloeon AcentrellaBaetiella and Liebebiella new genus (Ephemeroptera Baetidae)rsquo Journal of New YorkEntomological Society 95 553ndash568

Waltz RD and McCafferty WP (1989) lsquoNew species redescriptions and cladisticsof the genus Pseudocentroptiloides (Epherneroptera Baetidae)rsquo Journal of New YorkEntomological Society 97 151ndash158

Waltz RD McCafferty WP and Thomas A (1994) lsquoSystematics of Alainites n genDiphetor Indobaetis Nigrobaetis n stat and Takobia n stat (Ephemeroptera Baetidae)rsquoBulletin de la Societe drsquoHistoire Naturelle de Toulouse 130 33ndash36

Wang T-Q and McCafferty WP (1996) lsquoNew diagnostic characters for the mayfly familyBaetidae (Ephemeroptera)rsquo Entomological News 107 207ndash212

Weyenbergh H (1883) lsquoBijdrage tot de Kennis der zuid-amerikaansche EphemeridenrsquoTijdschrift voor Entomologie 26 159ndash174

Wiersema NA (2000) lsquoA new combination for two North American small minnow mayflies(Ephemeroptera Baetidae)rsquo Entomological News 111 140ndash142

Wiersema NA and McCafferty WP (2004) lsquoNew specific synonyms and records of NorthAmerican Centroptilum and Procloeon (Ephemeroptera Baetidae)rsquo Entomological News115 121ndash128

Wiersema NA Nelson CR and Kuehnl KF (2004) lsquoA new small minnow mayfly(Ephemeroptera Baetidae) from Utah USArsquo Entomological News 115 139ndash145

Wuillot J and Gillies MT (1993) lsquoCheleocloeon a new genus of Baetidae (Ephemeroptera)from West Africarsquo Revue drsquoHydrobiologie tropicale 26 213ndash217

Wuillot J and Gillies MT (1994) lsquoDicentroptilum a new genus of Mayflies (BaetidaeEphemeroptera) from Africarsquo Aquatic Insects 16 133ndash140

Zhou CF and Braasch D (2003) lsquoEine neue Gattung und Art der Heptageniidae ausdem ostlichen China (Ephemeroptera)rsquo Entomologische Nachrichten und Berichte 47147ndash151

Zhou CF Gui H and Su CR (2000) lsquoThe first record of the genus Brachycercus in Chinawith description of a new species (Ephemeroptera Caenidae)rsquo Entomologia Sinica 7132ndash134

Zhou CF and Peters JG (2003) lsquoThe nymph of Siphluriscus chinensis and additionalimaginal description A living mayfly with Jurassic origins (Siphluriscidae new familyEphemeroptera)rsquo Florida Entomologist 86 345ndash352

Zhou CF and Zheng LY (2001) lsquoRhithrogena trispina sp n a new species from China(Ephemeroptera Heptageniidae)rsquo Aquatic Insects 23 323ndash326

Zhou CF and Zheng LY (2003) lsquoTwo synonyms and one new species of the genusEphemera from China (Ephemeroptera Ephemeridae)rsquo Acta Zootaxonomica Sinica 28665ndash668

Zhou CF and Zheng LY (2004) lsquoThe genus Prosopistoma from China with description oftwo new species (Ephemeroptera Prosopistomatidae)rsquo Aquatic Insects 26 3ndash8

62 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13

Waltz RD and McCafferty WP (1987b) lsquoNew genera of Baetidae for some Nearcticpreviously included in Baetis Leach (Ephemeroptera)rsquo Annals of the Entomological Societyof America 80 667ndash670

Waltz RD and McCafferty WP (1987c) lsquoSystematics of Pseudocloeon AcentrellaBaetiella and Liebebiella new genus (Ephemeroptera Baetidae)rsquo Journal of New YorkEntomological Society 95 553ndash568

Waltz RD and McCafferty WP (1989) lsquoNew species redescriptions and cladisticsof the genus Pseudocentroptiloides (Epherneroptera Baetidae)rsquo Journal of New YorkEntomological Society 97 151ndash158

Waltz RD McCafferty WP and Thomas A (1994) lsquoSystematics of Alainites n genDiphetor Indobaetis Nigrobaetis n stat and Takobia n stat (Ephemeroptera Baetidae)rsquoBulletin de la Societe drsquoHistoire Naturelle de Toulouse 130 33ndash36

Wang T-Q and McCafferty WP (1996) lsquoNew diagnostic characters for the mayfly familyBaetidae (Ephemeroptera)rsquo Entomological News 107 207ndash212

Weyenbergh H (1883) lsquoBijdrage tot de Kennis der zuid-amerikaansche EphemeridenrsquoTijdschrift voor Entomologie 26 159ndash174

Wiersema NA (2000) lsquoA new combination for two North American small minnow mayflies(Ephemeroptera Baetidae)rsquo Entomological News 111 140ndash142

Wiersema NA and McCafferty WP (2004) lsquoNew specific synonyms and records of NorthAmerican Centroptilum and Procloeon (Ephemeroptera Baetidae)rsquo Entomological News115 121ndash128

Wiersema NA Nelson CR and Kuehnl KF (2004) lsquoA new small minnow mayfly(Ephemeroptera Baetidae) from Utah USArsquo Entomological News 115 139ndash145

Wuillot J and Gillies MT (1993) lsquoCheleocloeon a new genus of Baetidae (Ephemeroptera)from West Africarsquo Revue drsquoHydrobiologie tropicale 26 213ndash217

Wuillot J and Gillies MT (1994) lsquoDicentroptilum a new genus of Mayflies (BaetidaeEphemeroptera) from Africarsquo Aquatic Insects 16 133ndash140

Zhou CF and Braasch D (2003) lsquoEine neue Gattung und Art der Heptageniidae ausdem ostlichen China (Ephemeroptera)rsquo Entomologische Nachrichten und Berichte 47147ndash151

Zhou CF Gui H and Su CR (2000) lsquoThe first record of the genus Brachycercus in Chinawith description of a new species (Ephemeroptera Caenidae)rsquo Entomologia Sinica 7132ndash134

Zhou CF and Peters JG (2003) lsquoThe nymph of Siphluriscus chinensis and additionalimaginal description A living mayfly with Jurassic origins (Siphluriscidae new familyEphemeroptera)rsquo Florida Entomologist 86 345ndash352

Zhou CF and Zheng LY (2001) lsquoRhithrogena trispina sp n a new species from China(Ephemeroptera Heptageniidae)rsquo Aquatic Insects 23 323ndash326

Zhou CF and Zheng LY (2003) lsquoTwo synonyms and one new species of the genusEphemera from China (Ephemeroptera Ephemeridae)rsquo Acta Zootaxonomica Sinica 28665ndash668

Zhou CF and Zheng LY (2004) lsquoThe genus Prosopistoma from China with description oftwo new species (Ephemeroptera Prosopistomatidae)rsquo Aquatic Insects 26 3ndash8

62 J-L Gattolliat and C Nieto

Dow

nloa

ded

by [

Col

orad

o St

ate

Uni

vers

ity]

at 0

957

14

Sept

embe

r 20

13