Aquatic Entomology: Tour of the Orders Ephemeroptera (Mayflies) Ephemeroptera (Mayflies) Nymph D. Funk Imago - male D. Funk R4+5 MA1 MA2 MP1 MP2 Hind Wing Male gynadromorph (intersex form) of Neocloeon alamance Photo: D. Funk Mayfly Subimago/ Imago Flight Encalada, A.C. and B.L. Peckarsky. 2007. A comparative study of the costs of alternative mayfly oviposition behaviors. Behav Ecol Sociobiol 61:1437-1448. 1: Power-point presentation showing animations of oviposition behaviors of different mayflies (Ephemeroptera) observed in the upper East River drainage basin, near RMBL in western Colorado. A. Splashers (e.g., Siphlonurus occidentalis), B. Bombers (e.g., Drunella doddsi), C. Dippers (e.g., Rhithrogena hageni, Diphetor hageni, Epeorus longimanus), D. Landers (e.g., Baetis bicaudatus), E. Floaters (e.g., Acentrella turbinae). (To activate the animations, click on “ppt presentation” or the F5 key, and use the “return” key to move through the different animations (PPT 232 kb) Baetis Female Imago Ovipositing Examples of major aquatic habitat types in Maine, USA where mayflies occur. Mayfly nymphs are microhabitat specialists. Examples of Stream microhabitats Erosional with secondary depositional zones. Depositional with secondary erosional zones. Mayfly Mid-Network Maxima (MMM): RCC- Network Perspective Mayfly Diversity along Drainage Networks: MidNetwork Mayfly Maxima (MMM) diversity pattern predicted from overlap of species ranges along habitat gradient based on field observations. Large channel specialists replace small channel– headwater specialists, but overall diversity does not exceed mid-network maxima. A/In (Edge) Dorsal P/Out (Edge) Ventral Tibiopatellar Suture Ephemeroptera: Mayflies Caudal Filaments Abdominal Gills Prostheca (Planate) (Angulate) Galea-lacinia 3rd 2nd Maxillary Incisors (or “Canines”) Dentisetae (With or w/o serrations) Palpus Galea-lacinia Large apical setae with or w/o fine lateral branches or sometimes heavier serrations Baetis pluto Labium PG G Labial Palp 3rd 2nd 1st Mayfly Nymphal Diversity Arthropleidae Arthroplea sp. Baetiscidae Baetisca sp. Caenidae Ephemerellidae Sparbarus sp. Dannella sp. Behningiidae Dolania sp. Baetidae D. Funk Baetis tricaudatus Baetis bundyae Neocloeon triangulifer (2 gen., 2 species) Acanthametropus pecatonica (Alive – Wisconsin , May 2010, SK Burian) Characters [Nymph] – Prosternum with spine; claws longer than tibiae; labrum has wide median notch; maxillae has 2 canines, 2 well developed dentisetae, and rudimentary 3rd dentisetae Acanthametropus pecatonica -- Alive on native sand sediment from lower Wisconsin River, near Woodman, WI May 2010, SK Burian Ameletidae – (1 gen., 35 spp.) MP Fork D. Funk Ameletidae (A. lineatus) [By: D. Funk] Characters [Nymph] – Maxillae has apical pectinate setae; maxillae with canines reduced and only 1 dentisetae; mandibles elongate with reduced prostheca on R. man.; glossae of labium truncated with wide margin; labrum longer than wide. (1 gen., 2 spp.) (A. albrighti and A. neavei now jr. syn. of Ametropus fragilis) Ametropidae – (A. ammophilus) Characters [Nymph] – Foreleg reduced and palp-like with a distinct forcoxal projection; forclaw with setae; mid and hind claws long and sharp without setae; gills with long marginal setae. Ametropus neavei feeding behavior (Soluk and Craig 1988) [A. neavei now jr. syn. of Ametropus fragilis)] (27 gen., 157 spp.) Posterolateral projections absent Terminal filament small, but present small, but present Baetidae: Baetis sp. B. tricaudatus Characters [Nymph] – Epicranial suture lateral branches anterior to lateral ocelli; dorsal lobe of femoral apex directed ventrally; body similar in shape to ameletids and siphlonurids; hind wing pads present, absent, or minute; plate-like gills present on abd. Seg. 1-7, 1-5, or 2-7; some taxa have posterolateral projections on abd. seg. 9 and range from weakly to moderately developed (e.g., Callibaetis spp.) Baetidae (27 gen., 157 spp.) Acentrella (9 spp.) Americabaetis (3 spp.) Anafroptilum (8 spp.) Apobaetis (3 spp.) Baetis (21 spp.) Baetodes (16 spp.) Baetopus (1 spp.) Barbaetis (1 sp.) Callibaetis (11 spp.) Camelobaetidius (9 spp.) Caribaetis (1 sp.) Cloeodes (4 spp.) Cloeon (1 sp.) Diphetor (1 sp.) Fallceon (6 spp.) Heterocloeon (6 spp.) Iswaeon (3 spp.) Kirmaushenkreena (1 sp.) Labiobaetis (7 spp.) Moribaetis (1 sp.) Neocloeon (2 spp.) Paracloeodes (4 spp.) Plauditus (10 spp.) Procloeon (21 spp.) Pseudocentroptiloides (2 spp.) Varipes (1 sp.) Waynokiops (1 sp.) H. anoka H. curiosum Plauditus sp. villopore Labiobaetis sp. Antennal lobe Baetis tricaudatus Neocloeon triangulifer (formerly Centroptilum) Baetis bundyae D. Funk Acerpenna pygmaea (Female Nymph- NY) Acentrella rallatoma sp. n. (Male Nymph) A. macdunnoughi Plauditus dubius Plauditus virilis From Gary Lester Baetopus trishae Waltz from North Carolina (photo J. Webb) Blunt Apical Segment of Maxillary Palp Barbaetis benfieldi Kennedy (photo from J. Webb) Waynokiops dentatogriphus Photos: M. Bilger (2 gen., 9 spp.) Characters [Nymph] - Fore -claw bifid; tarsal claws of mid and hind legs single; maxillary palps with 2nd seg. Small; labial palps 2 segmented; body similar in shape to siphlonurids; body length 9-16 mm. By : D. Chandler Metretopus (2 sp.) Siphloplecton basale Siphloplecton ( 7 spp.) Siphlonuridae – (4 gen., 25 spp.) Siphlonurus sp. Siphlonisca aerodromia Siphlonuridae – (4 gen., 25 spp.) Edmundsius (1 sp.) Parameletus (4 sp..) Siphlonisca (1 sp.) Siphlonurus (19 spp) Characters [Nymph] – Sclerotized ridge (i.e., hind costa) of gills located medially not close to hind margin of gill lamellae; body fusiform, 6-20 mm; antennae short not longer than 2x’s the width of head; posterior abd. seg. with well developed posterolateral projections; 3 distinct caudal filaments; gills present on abd. seg. 1-7; labrum anterior margin smooth and sinuous. Medial Hind Margin Parameletus midas Parameletus midas Siphlonurus sp. By : D. Chandler ISONYCHIIDAE (1 gen., 16 spp.) Characters [Nymph] - Forecoxal gills filamentous; 1st pair of abd. gills dorsal in position; abd. gills have fibrilliform basal tuft. Isonychia obscura By : D. Chandler OLIGONEURIIDAE (2 gen., 9 spp.) Lachlania (4 spp.) Lachlania sp. Characters [Nymph] – Gills absent from leg bases; 1st pair of gills ventral, all other pairs of gills small, round, and dorsal; dorsal gills held close to surface of terga; galealacinia narrow apically; maxillary palp seg. 2 much longer than seg. 1; glossae fused into large labial plate. Homoeoneuria ammophila Homoeonuria (5 spp.) Homoeoneuria ammophila Filtering setae Characters [Nymph] – Tarsal claws subequal in length to tibiae; abd. gills lamellate with each lamella a basal tuft of filaments and a distinctive single, long, ventral appendage. P. centralis from Blackwater R., northern Florida – Jan Peters, FAMU (1 gen., 1 sp.) P. centralis from lower Wisconsin R., WI – May 2010, SK Burian P. centralis from lower Wisconsin R., WI – May 2010, SK Burian (Ametropus sp.) (Pseudiron centralis) [Chironomid prey buried in sand exposed by flow] (1 gen., 1 sp.) Characters [Nymph] – Maxillary palps extremely long extending posteriorly along the sides of the head Arthroplea bipunctata HEPTAGENIIDAE (14 gen., 126 spp.) By : D. Chandler Characters [Nymph] – Size 5-20mm; body dorsoventrally flattened; eyes positioned dorsally; epicranial portion of head capsule flattened and laterally expanded over the mouth parts; labial palpi with broad basal segment; abd. gills may be lateral or cupped beneath the body; gills present on segments 1-7 or 1-6 and most have a tuft of filaments at base; legs sprawling; 2 or 3 caudal filaments present. Heptageniidae (14 gen., 126 spp.) Afghanurus ( 13 spp.) Anepeorus ( 1 sp.) Cinygma ( 3 spp.) Cinygmula ( 10 spp.) Epeorus ( 19 spp.) Heptagenia ( 11 spp.) Ironodes ( 5 spp.) Leucrocuta ( 10 spp.) Macdunnoa ( 3 spp.) Raptoheptagenia (1 sp.) Rhithrogena ( 23 spp.) Spinadis (1 sp.) Stenacron ( 7 spp.) Stenonema ( 19 spp.) Heptageniidae: Heptagenia spp. H. pulla Cmp. eyes Cinygmula subaequalis By : D. Chandler Leucrocuta hebe Afghanurus lucidipennis (formerly genus Nixe) Leucrocuta sp Afghanurus spp. (Includes species formerly placed in genera Nixe and Ecdyonurus simplicioides species group) Dorsal Head (typical) Dorsal Abdominal Segments (typical) (11 gen., 87 spp.) Choroterpes basalis Characters [Nymph] - Body somewhat dorso-ventrally compressed, but not as great as in heptageniids; galea-lacinia with crown of long hair-like setae; hypopharynx well developed with well developed lateral lobes; maxillary palpi 3 seg.; At least 1 pectinate (i.e., comb or rake-like) spine present on inner anterior margin of maxillae; labial palpi 3 seg.; abdominal gills on seg. 1-7, 1-6, or 2-7 and some taxa may have gills on seg. 1 filamentous; most gills are bilamellate; 3 distinct caudal filaments Leptophlebiidae (11 gen., 87 spp.) Choroterpes (5 spp.) Farrodes (7 spp.) Habrophlebia (1 sp.) Habrophlebiodes (4 spp.) Hydrosmilodon (1 sp.) Leptophlebia (7 spp.) Neochoroterpes (4 spp.) Neoleptophlebia (7 spp.) Paraleptophlebia (32 spp.) Thraulodes (15 spp.) Traverella (6 spp.) Leptophlebia intermedia Leptophlebiidae: Neoleptophlebia assimilis All Gills Symmetrically Forked 1 7 Neoleptophlebia adoptiva Distinct Lateral Branches Paraleptophlebia moerens Deep fork - <1/4 Gill length Shallow fork - >1/3 Gill length Neoleptophlebia adoptiva BEHNINGIIDAE (1 gen., 1 sp.) Dolania americana Characters [Nymph] – Body ~13.0 mm; head with 2 distinct crowns of setae at anterolateral corners; tarsal claws absent on all legs, labial palpi long and stout resembling short prothoracic legs; ventrally oriented branched (i.e., forked) gills with edges finely divided (appearing fringed). Dolania americana (1 gen., 4 spp.) Characters [Nymph] – Frontal process of head absent; foretibiae round in cross section; abd. gill 1 single and filamentous; abd. gills 2-7 forked with margins extensively divided into filaments (appearing fringed); mandibular tusks usually curving in toward each other, but not up or down; mandibular tusks with scattered spines and stout setae; 3 distinct caudal filaments. Anthopotamus ( 4 spp.) Anthopotamus distinctus (4 gen., 7 spp.) Characters [Nymph] - Burrowing forms 12-35 mm; forlegs modified for digging; head with 10 internal air sacs; labial palpi not in the plane of the labium, but held beneath it and at a 90º angle to glossae and pararglossae; mandibular tusks curved in and downward apically; tusks with either upper or mesal surfaces with many tubercles; gills on abdomen held in dorsal position; abd. gill 1 vestigial; 3 caudal filaments. Ephoron leukon Polymitarcyidae (4 gen., 7 spp.) Campsurus ( 2 spp.) Ephoron ( 2 spp.) Tortopus (1 sp.) Tortopsis (2 spp.) (3 gen., 14 spp.) Hexagenia atrocaudata Characters [Nymph] – Body 12-32 mm and modified for burrowing; fossorial type legs (i.e., flattened and with edges specialized for digging); mandibular tusks curved upward at apex and outward; tusks also have a dorsolateral keel or ridge; tusks lack spines or have only basal spines; 3 caudal filaments. Litobrancha recurvata By : D. Chandler Ephemeridae (3 gen., 14 spp.) Ephemera ( 6 spp.) Hexagenia ( 7 spp.) Litobrancha (1 spp.) I attach a photo I took in northern Minnesota in spring of about 1957. I stopped my car for a few minutes on a bridge over a creek. I think that this was one of the 3 year and synchronized species. No longer possible to take this image. Daniel H. Janzen (1 gen.,1sp.) Characters [Nymph] - Mandibular tusks are longer than the head and with many long air-like setae arising from their entire surface; femora are flat an broad, but tibiae and tarsi are cylindrical and not apparently specialized for digging; gills on abdominal segment 1 are small and oval; gills on segments 2-6 are positioned laterally and have marginal filaments that makes them appear fringed and filaments are longer than the width of the lamella; antennae are much longer than head and lack setae; 3 caudal filaments have only inconspicuous setae Euthyplocia hecuba (1 gen., 1 sp.) Characters [Nymph] – Body 18-24 mm; mandibular tusks with short setae at tips and long setae at base; tusks crenate on outer upper edges; head with bifid process divergent apically; antennae lack setae; gills of abd. Seg. 1 forked to base; gill 2 dorsal lamella with outer edges of plate-like portion only slightly enlarged at base. Pentagenia vittigera Pentagenia vittigera (1 gen., 4 spp.) Characters [Nymph] – Gills on seg. 2 subquadrate, operculate, and fused along inner edge; other gills with many branches; maxillary palpi 3 segmented; labial palpi 3 segmented; hind wing pads present; eyes of male dioptic Neoephmera purpurea (7 gen., 37 spp.) Characters [Nymph] – No hind wing pads; interlocking or overlapping operculate gills on abdominal segment 2, but not fused; underlying gills lack branches. Caenidae (7 gen., 37 spp.) Amercaenis (2 spp.) Brachycercus (4 spp.) Caenis (13 spp.) Cercobrachys (7 spp.) Latineosus (1 sp.) Sparbarus (6 spp.) Susperatus (3 spp.) Caenis sp. Sparbarus maculatus (14 gen., 68 spp.) Gills Characters [Nymph] – Body subcylindrical to flattened, 5-15 mm; blunt spines or tubercles often on head, thorax and/or abdomen; abdominal segments usually have lateral projections; gills absent on abdominal segment 2; filamentous gills may be present on abdominal segment 1; lamellate gills present on segments 3-7 or 4-7 (operculate or semioperculate); 3 caudal filaments often with setae and spines. Penelomax septentrionalis By : D. Chandler Ephemerellidae: Drunella tuberculata Gills 3-7 7 3 Paired dorsal tubercles Fused wingpads Ephemerellidae (14 gen., 68 spp.) Attenella (4 spp.) Caudatella (5 spp.) Caurinella (1 spp.) Dannella (3 spp.) Drunella ( 12 spp.) Ephemerella (17 spp.) Eurylophella ( 16 spp.) Matriella (1 sp.) Penelomax (1 sp.) Serratella ( 5 spp.) Teloganopsis (1 sp.) Timpangoga (1 sp.) Tsalia (1 sp.) Ephemerellidae: Ephemerella spp. E. argo Semi- operculate Gills Tsalia berneri Paired dorsal tubercles Currently DNA barcode data indicate that Dentatella coxalis is deeply embedded in the Eurylophella and thus should be returned to that genus. LEPTOHYPHIDAE (4 gen., 35 spp.) Tricorythodes sp. By : D. Chandler Characters [Nymph] – Body small 3-10 mm; thorax stout in comparison with abdomen; abdominal segments 2-6 expanded; segments 7-10 not expanded; filamentous gills absent from abdominal segment 1; gills on abdominal segment 2 operculate oval or subtraingular; gills usually present on abdominal segments 2-6, rarely on 2-5; galea-lacinia narrow with setae restricted to crown or margin; labium with glossae very small or fused with paraglossae; labial palpi 2 or 3 segmented; hind wing pads present or absent. Leptohyphidae (4 gen., 35 spp.) Allenhyphes (1 sp.) Leptohyphes (11 spp.) Tricorythodes (22 spp.) Vacupernius (1 sp.) Tricorythodes sp. Tricorythodes explicatus - VT (1 gen., 11 species) Mesonotal Shield or “Carapace” Baetisca rubescens -- ME Characters [Nymph] – Body stout 4-14 mm; thorax with well developed mesonotum forming a carapace-like shield that extends to abdominal segment 6; abdominal gills are finely divided with the first pair covering succeeding pairs and all gills protected by enlarged mesonotum. Baetisca obesa – CT. River, CT [FFG – Gathering Collectors and Scrappers] Baetisca lacustris – CT. River, CT