Gripopterygidae Enderlein 1909

Key to larvae of South American Gripopterygidae

Family diagnosis. Gripopterygidae are distinguished by a supra ‐ anal retractable rosette of unsegmented, often purplish gill filaments. Only Notoperla lacks gills but can be separated from the gill ‐ less South American family, Notonemouridae, by its large furcasternum widely separating the coxae and by furcal pits located far from the spina. In Notonemouridae, the furcasternum is very small, coxae are close together, and furcal pits are adjacent to the spina.

44 46 47

The larva of Andiperlodes is unknown. Taxa that may or may not be dirt covered are identified by their morphology, not mainly by the dirt cover There is little information on morphological change during larval development of South American Gripo pterygidae, which may be important (Figs 43 ‐ 47). Change may also concern changes in late instars that are not related to wing development, for example Antarctoperlinae sp. II and IV. The key is to last instar larvae.

1 No gills; all coxae with an almost triangular projection next to articulation with thorax (Fig. 48 ‐ 49); large sprawling larvae with prominent hair fringes on legs; tibial spurs only in final instar (McLellan et al.2005) …………………………………………….. Notoperla spp.

1’ Anal gill rosette present, all coxa simple ……...…… 2

2 Dorsal face of abdomen with raised processes …….. 3

2 ʹ Dorsal face of abdomen at most with slightly elevated middle portion of rear margins but without raised processes, mostly completely simple ………………… 7

3 Abdominal processes unpaired ……………………… 4 3 ʹ Abdominal processes paired …………………………. 5

4 Sprawler habitus, distinct hair fringes on legs, gill rosette small; abdominal processes small, on some segments indistinct, thorax simple; abdominal sternites with dense cover of short erect hairs (friction pad?); lower face of all coxae entirely unsclerotised, a soft transparent membrane through which the muscles are clearly visible ………... Senzilloides panguipullii (Navás)

4 ʹ Perambulatory larvae, no distinct hair fringes; dorsal processes on abdomen and thorax very variable between species, and sometimes apparently also between sexes of a species; only eastern South America …………………………………………... Gripopteryx spp.

5 Head unarmed, anterior pronotal angles elongate, with pluridentate apex, posterior angles sharply pointed, raised; sometimes brachypterous, vestigial wingpads simple; abdominal processes smooth, simple, short, paraprocts ending in a strong downcurved spine, cercus short, base closely adjoined to dorsal paraproct edge ……………………… Neopentura semifusca Illies 1

5 ʹ Head with paired processes above antennal insertions or on vertex; other characters variable ……………… 6

6 Processes above bases of antennae are short simple cones; frontal depression anteriorly delineated by a horseshoe ‐ shaped blunt ridge; pronotum with sharply pointed extended corners, wing normal but ending in sharply pointed tip; abdominal processes small, simple; apex of paraproct narrow, tongue ‐ shaped …………… …………………………………… Ceratoperla fazi (Navás)

6 ʹ Processes on vertex, on all four corners of pronotum and on abdominal segments very long, narrow, spinose; wing pads with several marginal denticles and groups of setae; frontal depression very wide, no ridge in front of it; paraprocts sharply pointed ……………… …………….. Larventyp B (Araucanioperla?) of Illies, 1963

7 Prominent bulging eyes, no ocelli; gill very small ……. ………………………………… Andiperla willinki Aubert

7 ʹ Eyes normal, late instar larvae with at least rudimentary ocelli; large gill rosette ………………… 8

8 No tibial spurs................................................................ 9

8 ʹ Tibial spurs present; pleural membranes separate tergites from sternites on at least first three abdominal segments; cerci variable, often very long ………….. 20

9 Femora and tibiae with prominent caudal hair fringes; cerci slender with dorsal and ventral fringes of numerous hairs per segment; antennae long, bare; ocellar area of flat head with a depression; frons bulging distinctly forward above the clypeus on either side, next to the antennal foramen; labrum directed downward, at approximately right angle to flat top of head; maxillary palpus 5 ‐ segmented; tergite 10 caudally rounded; paraprocts short, truncate; slender larvae with rough integument; medium ‐ sized (12 ‐ 13 mm) ………………………….. Notoperlopsis femina Illies

9 ʹ No hair fringes; cerci short, less than half abdomen … …………………………………………………………..10

10 Basal abdominal segments with soft pleurae separating tergites from sternites; legs stout, femora with spinulose dorsal bulge; tarsus segments 1 and 2 subequal, barely longer than wide, 3 rd segment long, ventrally spinulose; tergite 10 sharply pointed; head wedge ‐ shaped in side view, with almost straight upper contour line; maxillary palpus 4 ‐ segmented, last segment long; a sharp raised cone near base of wingpads, similar smaller cones on pronotum; paraprocts long, narrow, upcurved (Larventyp X (Teutop.?) of Illies) …………………. Alfonsoperla gen. n.

1 Larventyp A (Araucanioperla?) of ILLIES, 1963.We agree with Vera (2006a) that Neopentura resembles, particularly in its male genitalia, the genus Zelandobius (McLellan, 1993) from New Zealand, more than other South American genera. However it differs from Zelandobius by its peculiarly forked male paraprocts and the appendage on the tip of the female subanal lobes. We have seen rich material, including adults, from several sites in Neuquen, Argentina.

10’ Abdominal tergites and sternites forming continuous rings, no abdominal pleurites (Fig. 51); legs normal, femora simple, tarsi slender, basal segment much longer than wide and distinctly longer than 2 nd, 3 rd segment not spinulose; remaining Antarctoperlinae … ………………………………………………………….. 11

11 Sides of meso ‐ and metanotum flanged; flanges have kinks and extensions mimicking early wing pads; they are bifid on metathorax; apex of paraproct blunt; cerci stout, shorter than last segment, basally thick, conically tapered, curved upward; large [up to 25 mm long], wingless; terrestrial larva (Illies 1960)........................... ................................................ Megandiperla kuscheli Illies

11 ʹ Sides of meso ‐ and metanotum not flanged, wingpads normal, sometimes with sharply pointed apex; cerci thin, neither conical nor downcurved ……………... 12

12 Heavy detrital coating concealing body contour, except strikingly clean segment 10; dorsal surface without ground pilosity, bare except for localized patches or lines of long setae; base of antennal flagellum much thickened, with dense cover of long curled hairs …. 13

12 ʹ Body contour remains clearly visible even if some dirt is trapped in variable pilosity; segment 10 not strikingly cleaner than others; base of antennae not much thickened but may be densely pilose ……….. 14

13 Front corners of pronotum anterolaterally projecting; groups of long setae on front corners of pronotum, shoulders of mesothorax, tips of hind wing pads, and on abdominal tergites 1 ‐ 9 which lack regular setal fringes ……………………………………………………. ……... “ Larval type C (Araucanioperla?)” of Zwick, 1972

13 ʹ All four corners of pronotum rounded; long hairs in rows along outer edges of thorax (including wing pads) and caudally on abdominal tergites 1 ‐ 9 ……………….. ……………………………. Pelurgoperla personata Illies 2

14 Paraprocts broad, apex rounded; fringes of straight setae around perimeter of pronotum, a few along shoulders, and along distal margins of abdominal segments; ventral face with soft, often curly hairs; pilosity may trap some mud ………... Chilenoperla spp.

14 ʹ Paraprocts terminate in a point; point varies in length …………………………………………………………… 15

15 Paraprocts obliquely truncate, medial tip short ……… …………………………………… Antarctoperla spp..... 16 15 ʹ Paraprocts with prolonged distinctly offset spine … 17

16 Dorsal side with many long, curly (not straight) hairs which may trap dirt; ventral face with pale short very dense hair cover; tergite X with long dorsolateral setae and a short apical setal fringe (Figs. 50 ‐ 52). Males with shallow posteromedial keel on tergite 10; pronotal corners rounded …………… Antarctoperla altera Zwick

2 Hairs on antenna are curved, not straight as in fig. 43b of Illies (1963).

16 ʹ Dorsal side without dense long pilosity; pronotal corners a little produced but blunt; tergal fringes consist of blunt or club ‐ shaped pale setae shorter than intersegmental membranes; paramedian brown bands lengthwise on pronotum and over wingpads, a median brown band along middle of abdomen ………………......................................... Antarctoperla michaelseni (Klap.)

17 Rear pronotal corners rectangular, not projecting or even horn ‐ like; no long hairs on body ……………... 18

17 ʹ Rear pronotal corners prominent, sometimes resembling short horns; at least some long hairs on body …………………………………………………… 19

18 Stout larva; sides of pronotum straight, anteriorly slightly converging, corners rounded; paraproct truncate, medial edge extended into long, almost straight spine; regular fringes of short dark blunt setae along caudal edges of abdominal segments; base of antennal flagellum somewhat thickened, densely setose; present specimens short ‐ winged but may be penultimate instar (Figs. 24 ‐ 29) ………………………................................................. Antarctoperlinae gen. sp. II

18 ʹ Slender larva; sides of pronotum parallel and slightly concave. Front pronotal corners a little extended; abdominal fringes of slender pale hairs with tips curved against body; paraproct caudally narrowed, obliquely truncate, inner edge blending into a long, upcurved spine; micropterous (Figs. 30 ‐ 36) …………................................................ Antarctoperlinae gen. sp. III

19 Wing pads normal, not flanged; abdominal fringes composed of inconspicuous short spatulate setae; contour of abdominal segments in dorsal view sinuous; pronotal horns pointing sideways in immature larvae, slightly convergent in last instar; 3 dark rings on antennae, each about 3 segments long; caudal margins of abdominal tergites with short spatulate setae and long interspersed hairs. A dark narrow median line from rear margin of pronotum to tip of abdomen; paraproct with short sharp dark point (Figs. 37 ‐ 43) …............................................. Antarctoperlinae gen. sp. IV

19 ʹ Wing pads flanged, very wide; pronotal horns slightly divergent, apparently a little variable with instar; erect bristles along edges of abdominal tergites; contour of abdominal segments in dorsal view strongly bell ‐ shaped, widening from narrow base in a convex curve, then again constricted in front of even wider, bulging distal edge; no dark longitudinal line; a dark very sharp point on paraproct (Figs. 18 ‐ 23) …… ……………………………… Antarctoperlinae gen. sp. I

20 Hind margin of metanotum and also mesonotum re entrant, angularly excised (Fig. 54) …………………… ……………………………….. Limnoperla jaffueli (Navás) Slender larva, cerci about half as long as abdomen. Body pilosity very fine, barely visible. Sparse setation on dorsal side of femora somewhat irregular (Fig. 64). Tibial hair fringe fine, sparse.

20 ʹ Hind margin of thoracic nota straight to slightly convex (Fig. 53); larvae of various shapes ………………….. 21

21 Slender, very hairy larvae; long, soft, pale pilosity may trap dirt; antenna and legs with prominent setal fringes, also along upper and lower edges of cerci ….. …………………………. Potamoperla myrmidon (Mabille)

21 ʹ Body lacks apparent pilosity; hair fringes, if present, sparse, no long hairs on antenna …………………… 22

22 Paraproct tongue ‐ shaped, wide, in ventral view concealing basis of cercus; stout larvae …………….. 23

22 ʹ Paraproct of various shapes, not laterally expanded, cercus base in ventral view exposed ……………….. 24

23 Femora simple, slender (length/width ~ 3.7); an elongate bulge at wingbase; large triangular apophyses on front margin of tergite X are easily seen by transparency………...….. Aubertoperla illiesi (Froehlich) Tiny stout peg ‐ like setae on occiput, near midline and anterior margin of meso ‐ and metanotum and along distal edges of abdominal tergites; sparse elsewhere, absent from head in front of occipital suture and from antennae and legs. Body surface in addition to pegs with fine, indistinct pilosity, hairs with wide bases and narrow long apices. Paraprocts short and wide, in ventral view covering the cercus base. Cerci long, segment 5 or 6 distinctly longer than wide.

23 ʹ Femora wide and flattened (length/width ~ 3.2), trian gular in cross section, the fore femur in particular, posterodorsal edge crest ‐ like; no distinct bulge at wingbase; tergite X without anterior apophyses; integument densely covered with only short peg ‐ like setae, no interspersed slender hairs …………………… ………………………….…...… Aubertoperla kuscheli Illies

24 Each femur with a (sometimes small) ventral spine; no hair fringes on legs and antennae; only eastern South America ……………………………………………….. 25

62 63 64

24 ʹ Femora structurally simple; hair fringes present or absent …………………………………………………. 26

25 Slender narrow larvae of normal habitus, with long antennae and legs ………………… Tupiperla Froehlich

25 ʹ Stout larvae, thorax much widened, head set between anteriorly extended pronotal corners …………………. ………………………………….. Guaranyperla Froehlich

26 Stout larvae; body, perimeter of pronotum, scape, pedicel, and legs covered with small spine ‐ or peg ‐ like setae …………………... Paragripopteryx Enderlein spp. We studied P. anga Froehlich, P. blanda Froehlich, P. klapaleki Enderlein, and P. munoai Benedetto.

26 ʹ Slender larvae ………………………………………... 27

27 Aculeate setae cover body and edges of nota; wingless …………………………. Falklandoperla kelper McLellan Falkland Islands only

27 ʹ No setation along edge of pronotum, hairs on body minute, often only visible under a compound microscope …………………………………………… 28

28 Maxillary palpus 4 ‐ segmented, with long last segment; legs without hair fringes; cerci glabrous except for short apical whorls of setae (Figs. 55 ‐ 57) ……………… …………………………... Claudioperla tigrina (Klapálek) Pronotal and mesonotal pilosity (requires high magnification!) consists of setae with oval base tapering abruptly towards a much longer, filiform distal section. Setation on anterior abdominal segments similar, difference between wide base and flagellum ‐ like apex of setae becomes successively less pronounced on posterior abdominal segments. Setae on TX almost unmodified. Intersegmental membrane with numerous floriform chloride cells on long pedicels. Integument finely shagreened, with isolated asperities.

28 ʹ Maxillary palpus 5 ‐ segmented; hair fringes on legs present but not obvious; one dorsal and one ventral seta in each apical whorl on cercus segments long, forming regular sparse vertical fringes …………… 29

29 Frontoclypeal suture medially interrupted, frons forms little lateral bulges (Fig. 61); femur and tibia with single regular row of erect hairs; femur with very few minute spines along ventral edge (Fig. 62); marginal setae of abdominal segments very short, truncate, spatulate, very different from hairs on tergite; no chloride cells along tergal intersegmental membranes, the few spotted on thoracic membranes with much shorter pedicle ……………………………………... Rhithroperla (our material includes a mature male of R. rossi)

29 ʹ Frontoclypeal suture entire, clypeus completely separate from frons; hair fringe on femur plurilinear, dense, some short spines to outside of fringe (Fig. 60) also present, also on tibia; body hairs with thickened base and narrow apex, marginal setae a little larger but otherwise similar; shagreen consists of sparse isolated cuticular asperities; numerous chloride cells on tergal intersegmental membrane, pedicle shorter than in Claudioperla …………………………………... Teutoperla We studied pharate males of T. rothi Illies, 1963 and T. brundini Illies, 1963.