The Leptophlebiidae: Atalophlebiinae of New Caledonia (Ephemeroptera). Part VIII: systematics

– Neampia , a new genus of Atalophlebiinae (Leptophlebiidae) from New Caledonia, is described for the type species Neampia quaternaria new species, and ﬁve other new species. Of the three new subgenera established, three new species, Neampia quaternaria , Neampia inspina and Neampia richardi , belong to Neampia s.s . The second new subgenus, Goa , is monotypic and contains only the new species Neampia natans . The third new subgenus, Anodela , contains two new species, Neampia ﬂuvialis and Neampia septentrionalis . Four of the new species are directly associated with nymphs by rearing, and two are associated by dissection of the developing genitalia in mature male nymphs.


Introduction
This paper is the eighth in a series on the systematics, phylogeny and biogeography of the Leptophlebiidae (Ephemeroptera) of New Caledonia.Part I of this series lists all localities (Peters et al., 1978).Other works are Peters and Peters (1980, 1981a, 1981b, 2000) and Peters et al. (1990Peters et al. ( , 1994)).These publications and original localities and data of Starmu¨hlner (1968) are available on Ephemeroptera Galactica (http://www.ephemeropteragalactica.com).For convenience, localities are repeated in Table Al and source of material for figures in Table A2.Neampia, a new genus with three subgenera (Neampia s.s., Goa and Anodela new subgenera) is described here.The genus is characterized by males with long, narrow penes with apical spines and females with an egg guide composed of an apical elongation of sternum 7 joined with a sclerotized anterior prolongation of sternum 8.The nymphs are small with compact bodies, short antennae, narrow gills and apically divergent clypeal margins (Figs. 42,(43)(44)(45)(46)(47)(48)(49)(50).
Methods are those described in previous sections."Mature nymphs" refer to nymphs with well-developed wing pads -not necessarily last instar nymphs.In the male forelegs, the lengths of successive segments are compared with the length of the tibia and expressed as a ratio; the length of the tibia is given in parentheses.
Repositories for the type specimens are abbreviated as: Florida A&M University, Tallahassee, Florida, USA (FAMU), an affiliate of the Florida State Collection of Arthropods; Bernice P. Bishop Museum, Honolulu, Hawaii, USA (BPBM); National Museum of Natural History, Washington, DC, USA (NMNH); and the Muse´e cantonal de zoologie, Lausanne, Switzerland (MZL).
Egg (description based on N. (Goa) natans).Length and width 145r 70 mm 2 ; covered with circular structures,  Genitalia,ventral view (17,21,24,26,29,33),lateral view (20,23,28,30);detail,apex of penes,25,27,[31][32]dorsal (22).Figure 33 55) or curved with shallow depression distal to curve.Lingua of hypopharynx with well-developed lateral processes, anterior margin cleft, apex of submedian lobes with rake-like process; superlinguae as in Figure 56, with row of setae along anterior margin, lateral margins blunt.Segment 2 of maxillary palpi subequal in length to segment 1; segment 3 of palpi more than two-thirds to slightly shorter than length of segment 2, triangular; V-shaped ridge near ventral inner anterolateral margin of maxillae; setae on maxillae as in Figure 59.Labium as in Figure 57; segment 2 of palpi slightly shorter to equal length of segment 1; segment 3 of palpi three-fifths to fourfifths length of segment 2; paraglossae ventral to glossae.-9 (34, 37); 4-6 (38, 40-41); female subimago terga 3-7 (35); 36, lateral abdominal segments 5-6 (36), 5 (39).(Figs. 44 and 45); small unbranched tracheal branches occasionally present basally (Fig. 45); main trunk and branches pigmented, both portions of lamellae hyaline or grayish translucent.Posterolateral spines on abdominal segments 2 to 5-9, spines progressively larger posteriorly, apex of posterolateral spines on segments 9 or 8-9 indented and giving appearance of double spines.Terminal filament is slightly longer than cerci; caudal filaments with whorl of spines and setae of moderate length at articulations.middle of anterior margin (Figs.3-6); (5) claws of a pair are dissimilar; one is apically hooked and the other is obtuse, pad-like (Fig. 11); (6) penes are narrow, tubular and fused except at apex (Figs.17-33) and each lobe usually possesses a small apical spine; and (7) female has a well-developed egg guide which is composed of the posterior projection of sternum 7 and a sclerotized anterior projection of sternum 8 (Figs.14-16).In the nymph: (1) anterior margin of lingua of hypopharynx is cleft and inner apex of each submedian lobe has a rake-like Within New Caledonia, Neampia appears most closely related to Amoa Peters & Peters from which it is distinguished in imagos by the narrow and tubular penes of the male and the well-developed egg guide of the female.The nymph can be distinguished by the divergent margins of the clypeus, the generally compact body shape and (in subgenus Goa) by the highly modified femora; also, the developing egg guide is usually visible in mature female nymphs.The short antennae (< 2 times length of head) of Neampia are a useful character for distinguishing mature nymphs, but antennae are longer in young nymphs.
Discussion.The characters above will distinguish the male imagos and nymphs of Neampia (s.s.) from those of the new subgenera Goa and Anodela.A single female subimago is designated as allotype of the type species Neampia (s.s.) quaternaria because the specimen is reared and intact hind tibiae are present on the exuviae to confirm the association.The apex of the 9th sternum of this female is truncate, but it is rounded on smaller females from other localities, such as Neampia sp. (Fig. 13) collected with males of N. quaternaria at N50.This difference may be size-dependent rather than a character of species.All females of Neampia (s.s.) appear to have the same general color pattern (Fig. 35), and unassociated females cannot be placed to species.The egg guide of these females is similar to those illustrated (Figs. 14 and 15).The egg guide of subimagos is not as expanded as that of imagos.Female imagos and subimagos of Neampia (s.s.) spp.are recorded from N27, N35, N42, N50 and N54.
Male subimago (in alcohol).As in male imago, except for usual developmental differences; wing membrane tan.
Female imago.Unknown.Female subimago (in alcohol).Length: body, 5.4 mm; forewings, 6.2 mm.Eyes black.Head light brown, carinae darker, washed with scattered blackish markings, sutures lighter.Antennal scape and pedicel dark brown, flagellum (broken off and missing) presumably paler as in male.Thorax: dark brown, carinae and median longitudinal suture darker, sutures paler, carinae of pleura and pronotum extensively washed with black.Markings of legs as in male imago, except base color of legs light brown.Wings: color and marks as in male imago, except longitudinal veins of fore and hind wings darker brown and membrane light tan.Abdomen: light brown, terga 2-6 with dark brown, posterior band on each tergum, tergum 1 uniformly washed with dark brown, tergum 10 uniformly washed with brown, terga 3-7 with color pattern as in Fig. 35; spiracles dark brown, tracheae hyaline; sterna 2-6 with light brown, lateral stripes, sterna 8 and 9 uniformly washed with darker brown; egg guide weakly developed, washed with blackish brown.Caudal filaments light brown, paler at base, articulations pale.
Mature nymph (in alcohol).Head: light brown, washed with brown anterior to eyes and between ocelli, clypeus and area anterior to median ocellus pale.Labrum with concave emargination and five small, rounded denticles (Fig. 46).Thorax brown, venter lighter, lateral areas of mesonotum, median longitudinal suture, posterior portion of scutellum and areas around forewing pads washed heavily with darker blackish brown; pronotum with submedian transverse bars and lateral margins darker brown.Legs: pale; mid and hind coxae with brown macula; apex of femora with small dark brown macula; tarsi darker brown; outer margin of hind tibia with row of 27-40 closely spaced, small, spine-like setae (Fig. 67).Abdomen: color and marks as in male and female imagos.Gills (Fig. 44): membrane hyaline, tracheae black.Caudal filaments pale.
Etymology.Quaternarius, L., meaning consisting of four, in reference to pale abdominal color pattern on terga 2-5 of males.
Diagnosis.Neampia (s.s.) quaternaria can be distinguished from the remaining species of Neampia (s.s.) by the following combination of characters.In the imago: (1) abdominal terga 1 and 6-10 of male are opaque, dark brown to lighter brown posteriorly, and terga 2-5 are hyaline, washed with light brown (Fig. 34); and (2) apex of each penis lobe has a small apical spine and the openings of penes are dorsoapical .In the nymph: (1) abdominal terga 1 and 6-10 of the male are dark brown and terga 2-5 are pale (as in Fig. 34); (2) anteromedian emargination of the labrum is concave; (3) clypeus is pale; and (4) outer margin of hind tibia possesses a row of closely spaced, small, spine-like setae (Fig. 67).
Discussion.The size range given for the males includes two specimens from N37, a small dark brown male subimago (body length 3.6 mm) and a large male imago (body length 6.0 mm, wing 6.4 mm).For all other material, the body size of males ranges from 4.0 to 5.0 mm.Specimens from the southern serpentine area are small and generally darker in color at localities N53 and N50, but slightly lighter at N51 (darkest on tergum 6 and apical half of tergum 7 and slightly paler on terga 8-9); at N54, the pattern is lighter overall and the blackish wash on tergum 6 may be less intense than that illustrated.The abdominal color pattern of female nymphs is variable throughout the range.Posterolateral projections always occur on terga 5-9, but smaller posterolateral projections on terga 3-4 or 4 may also be present.This character, like the number of spine-like setae on the hind tibiae, can vary in individuals between the left and right side of the body.The single female subimago assigned to this species is discussed under the subgenus.
Biology.Neampia (s.s.) quaternaria occurs throughout New Caledonia and was found in small streams to large rivers at altitudes of 6-458 m and water temperatures of 16.5-23 xC, but were most abundant at temperatures of 18-20.5 xC.Nymphs were found under small rocks along the river banks in slow moving water.Subimagos were collected at light traps.
Neampia (Neampia) inspina, new species (Figs.21-23, 47 and 68) Male imago (in alcohol).Length: body, 4.0-4.8mm; forewings, 4.8-5.2mm.Upper portion of eyes reddish brown, lower portion black.Head dark brown, carinae darker.Antennal scape and pedicel dark brown, flagellum light brown.Thorax dark brown, carinae darker, sutures paler, carinae of pleura and pronotum washed heavily with black.Coxae washed with brown, forefemora light brown, remainder of legs pale with apical margins of foretibiae and femora of all legs with brownish markings, apex of tarsal segments brownish.Wings: longitudinal veins light brown basally, slightly darker in apical fourth of cells C, Sc, and R1 of forewings; membrane of fore and hind wings hyaline, except base of fore and hind wings brown, cells C and Sc of forewings translucent.Abdomen: light brown, terga 1-9 with dark brown, wide, transverse band on posterior margin of each tergum, tergum 1 uniformly washed with dark brown, terga 2 and 3 washed heavily with dark brown except for narrow median area and lateral margins, terga 4 and 5 pale except for enlarged posterolateral blackish brown maculae above spiracles; terga 6-10 washed with dark brown similar to Figure 34; spiracles dark brown, tracheae washed lightly with dark brown; sterna 1-6 light brown, sterna 7-9 lightly washed with brown.Genitalia (Figs. 21-23): brown, apical half of forceps and penes paler; penes with dorsal openings, but without spines.Caudal filaments pale with dark articulations in basal half.
Material.Holotype male imago, No. N54; paratypes: one nymph, No. N13; two nymphs, No. N15; one nymph, two males, No. N35; six nymphs, No. N53; eight nymphs, one male, No. N54.All types are in alcohol.Association of the nymphs and adults is by dissection of developing male genitalia from nymphs from the same locality.Types are deposited in the following collections: holotype, one male and eight nymphal paratypes at FAMU; one male and four nymphal paratypes each at MZL and NMNH, and two nymphal paratypes at BPBM.Other specimens appearing to belong to this species include nymphs from N27 and FNK53.
Discussion.All imagos are from a single locality on two dates (N35, N54) and associated nymphs have a light brown clypeus with few markings.The clypeus on associated nymphs from N53 is also light brown but with darker brown apical and lateral markings.The color pattern of male imagos is similar to that of N. richardi n. sp.from which it is distinguished by the lack of spines on the penes.
Biology.Neampia (s.s.) inspina has been collected on the West Coast from Nondoue´to Paı¨ta and from tributaries of the Rivie`re Bleue.The species is found in large streams at 31-122 m with water temperatures of 18-20.5 xC.Adults were collected as subimagos at light traps and kept until they transformed to imagos.
Discussion.The brown coloration on terga 2-4 is more faded in the male from N40.The color pattern of male imagos is similar to that of N. inspina n. sp.from which it is distinguished by the presence of spines on the penes.
Biology.Neampia (s.s.) richardi was collected from small streams to large rivers on the East Coast and in the southern region at altitudes from 31 to 183 m and water temperatures of 19-24 xC (most abundant at water temperatures of 19-20 xC).Subimagos were collected at light.Little is known about the biology of the species.
Etymology.Goa, based on the name of the town Goa, New Caledonia.Feminine.
Type species.Neampia (Goa) natans new species.Discussion.The characters listed above will distinguish the imagos and nymphs of Goa from those of Neampia (s.s.) and Anodela.
Male subimago.Markings as in male imago; wing membrane light brown.
Female imago (in alcohol).Length: body 5.5-6.3mm, forewings 7.0-7.1 mm.Eyes black.Head dark brown, carinae darker, sutures pale.Antennal scape and pedicel dark brown, flagellum lighter.Thorax: color and marks as in male imago.Coxae dark brown, remainder of legs brown to light brown, apex of femora dark brown, foretibiae with narrow, dark brown, transverse distal band, mid and hind tibiae darker at apex, foretarsi and tarsal segments 4 of mid and hind tarsi washed with dark brown.Wings: longitudinal veins of fore and hind wings dark brown, cross veins of fore and hind wings brown, except those in anterior half of forewings slightly darker, membrane of fore and hind wings light brown, except base of fore and hind wings darker, cells C and Sc of forewings translucent.Abdomen: color and marks as in male imago, except general color darker, sublateral bars on terga 4 and 5 larger, and sterna 2-9 brown with lateral margins washed lightly with darker brown.Egg guide washed lightly with dark brown, posterior portion darker brown (Fig. 14).Caudal filaments pale.
Female subimago.As in female imago, except egg guide less developed and wing membrane light brown.
Mature nymph (in alcohol).Head: light brown, washed with darker brown anteriorly, between ocelli and on dorsum as in Figure 1.Thorax: dark brown, venter pale, pronotum pale with darker blackish-brown marks; lateral areas and posterior margin of forewing pads washed heavily with darker blackish brown (Fig. 1).Legs: pale, mid and hind coxae with dark brown macula, apex of femora dark brown, a larger dark brown, mid-dorsal macula on femora as in Figures 1 and 61, marks on tibiae as in male and female imagos, except darker, tarsi dark brown.Abdomen (Fig. 1): color and marks as in male and female imagos, except terga 8-10 paler, sterna uniformly pale except anterior margin of sterna 8 and 9 washed lightly with darker brown.Gills (Fig. 45): membrane hyaline, tracheae black.Caudal filaments pale.
Diagnosis.Neampia (Goa) natans sp.n. is the only species known in this subgenus and is recognized by distinctive characters of the subgenus.
Discussion.Males are generally smaller than females.The color pattern is less intense in some specimens and leg markings are darker and more pronounced in females.
Biology.Neampia (Goa) natans has been collected from a few mountain streams along the East Coast but most were found at one undisturbed site at a higher altitude (458 m).Water temperatures were 18-19 xC when collected (15.2 xC at FNK79).Nymphs were excellent swimmers.They were found in still pools along the banks and at the bottom of waterfalls resting on rocks, but when disturbed they swam quickly to another part of the pool (usually under edges of rocks).As quick, darting swimmers, their behavior resembles that of members of the Baetidae.
Nymphs apparently prefer very clean water.Where a dirt road forded the stream (locality No. N37), nymphs were observed above but not below the ford although there was only a slight increase of silt.Most other species occurred both above and below the ford.
Subimagos emerged from 1500 to 1900 h with most emerging before sunset (1800 h).Imagos molted the next morning.Swarming was not observed, and no specimens were collected at light.
Etymology.Anodela, an arbitrary combination of letters, partially based on anagram of Caledonia.
Type species.Neampia (Anodela) fluvialis, new species.Discussion.The characters above will distinguish the imagos and nymphs of Anodela from those of Neampia (s.s.) and Goa.A few other unassociated nymphs of Neampia (Anodela) collected at various localities in the southern region and along the West Coast are not assigned to species.
Male subimago (in alcohol).As in male imago, except for usual developmental differences; wing membrane grayish tan.
Female imago (in alcohol).Length: body 5.5 mm; forewings 5.5 mm.Color and markings as in male imago, except mesothorax lighter brown.Margin of sternum 7 pale, anterior margin of sternum 8 shaded dark gray.
Female subimago (in alcohol).Color and markings as in male imago; wing membrane grayish tan; egg guide weakly indicated, not extended as in imagos.
Mature nymph (in alcohol).Head: brown, venter lighter, dorsum washed heavily with darker blackish brown between eyes and ocelli (Fig. 42), clypeus pale.Thorax: dark brown, venter lighter, lateral area of mesonotum and areas around forewing pads washed heavily with dark blackish brown, pronotum with broad, blackish-brown median macula as in Figure 42.Legs: light brown; mid and hind coxae with dark brown macula; femora washed lightly with dark brown, darker apically, tarsi reddish brown (Fig. 62).Abdomen: color and marks as in male imago, except median longitudinal line on terga light brown.Gills (Fig. 42): membrane gray, tracheae black.Caudal filaments brown.
Etymology.fluvialis, L., referring to a river, in this case the type locality Rivie`re Bleue.
Diagnosis.Neampia (Anodela) fluvialis can be distinguished from the other species of Neampia (Anodela) by the following combination of characters.In the imago: abdominal terga 1-10 of the male are dark brown and heavily washed with blackish brown (Figs.38 and 40), and (2) apex of each penis lobe of male genitalia has a strong, ventral spine .The same color pattern applies to the nymphs: (1) abdominal terga 1-10 are dark brown (Fig. 42).
Discussion.There are several small population variations not included in the type series; these might eventually represent additional species or subspecies but are here treated as variations of N. (A.) fluvialis.In imagos from the paratype series, tergum 5 is either dark brown (as the rest of terga) or possesses a median macula only a shade grayer than that on other terga (Fig. 40), and femora are light brown basally and darker apically.Variations include: 1. Femora blackish (two nymphs, N25; two male imagos and one male subimago, N53). 2. Posterior portion of tergum 5 pale without blackish submedian longitudinal line (nymphs only, N27, N55). 3. Three nymphs from N37 seem to have the color pattern of N. (A.) fluvialis, but all other specimens from these localities belong to N. (A.) septentrionalis n. sp.In the absence of male imagos, these specimens are treated here as a variant.
Biology.The type series of Neampia (Anodela) fluvialis was found in the southern region and in medium to largesized rivers.Nymphs were found in streams with water temperatures of 19-20.5 xC at elevations from 150 to 183 m.The species was most abundant around the Rivie`re Bleue.Nymphs were collected under small rocks along the river bank in slow moving water.Subimagos were collected at light, but swarming was never seen.Neampia (Anodela) septentrionalis, new species (Figs. 6,15,33 and 41) Male imago (in alcohol).Length: body, 4.5 mm; forewings, 4.5 mm.Upper portion of eyes reddish brown, lower portion black.Head dark brown, carinae darker.Antennal scape and pedicel dark brown, flagellum much paler.Thorax dark brown, carinae darker, sutures paler, carinae around bases of legs and wings washed heavily with darker brown, margins and carinae of pronotum washed heavily with darker brown.Coxae dark brown, remainder of legs pale, except forefemora slightly darker, apex of femora and tibiae dark brown.Wings: longitudinal veins of fore and hind wings light brown, cross veins hyaline except those in cells C of forewings pale brown; membrane hyaline, apical third of cells C and Sc of forewings translucent.Abdomen: light brown, terga 1-10 washed with blackish brown except for large pale macula on tergum 5, terga 6-10 darker brown, terga 1-10 with narrow, pale, median longitudinal line extended entire length of each tergum (Fig. 41) and with paired pale sublateral triangular-shaped marks next to submedian blackish lines on terga 2-8 (Fig. 41); spiracles dark brown, tracheae hyaline; sterna 8 and 9 uniformly washed with darker brown.Genitalia and caudal filaments broken off and missing.
Male subimago (in alcohol).As in male imago, except for usual developmental differences; wing membrane light brown.Genitalia as in Figure 33.
Female imago (in alcohol).Length: body 4.0-4.8mm; forewings 5.2-5.5 mm.Eyes black.Head brown, area between eyes and ocelli darker.Antennal scape and pedicel brown, flagellum paler.Thorax: color and marks as in male imago.Wings: longitudinal veins in fore and hind wings brown, cross veins light brown, except faded in posterior half of wings; membrane of fore and hind wings hyaline to light brown, except base of wings darker, apical third of cells C and Sc of forewings translucent.Color and marks of legs as in male imago, except tarsi washed lightly with darker brown.Abdomen: color and marks as in male imago, except pale submedian marks on terga absent, lateral margins of sterna 1-7 washed lightly with darker brown.Egg guide as in Figure 15.Caudal filaments brown.
Mature nymph (in alcohol).Head: brown, venter lighter, dorsum washed heavily with darker brown between eyes and ocelli.Thorax: dark brown, venter lighter, lateral areas of mesonotum and areas around forewing pads washed heavily with darker brown.Pronotum with darker brown median macula similar to that in Figure 42.Legs: pale, mid and hind coxae with brown maculae, other color and marks as in male and female imagos, except tarsi reddish brown.Abdomen: color and marks as in imagos.Gills: membrane gray, tracheae blackish gray.Caudal filaments brown.
Diagnosis.Neampia (Anodela) septentrionalis can be distinguished from the other species of Neampia (Anodela) by the following combination of characters.In the imago: abdominal terga 1-10 of male are brown, heavily washed with dark blackish brown and with a large pale macula on tergum 5 (Fig. 41).The same color pattern character applies to nymphs: (1) abdominal terga 1-10 dark brown with a large pale macula on tergum 5.
Discussion.The intensity of the coloration on the egg guide is pale in some specimens.
Biology.Neampia (Anodela) septentrionalis occurs along the East Coast and is found in small to large streams.Nymphs were collected in streams with water temperatures of 15 -20 xC and elevations of about 20-458 m.
, adults of Neampia will key to Tepakia Towns & Peters, from which they are distinguished by the dissimilar claws, an unforked MP vein in the hind wing, and the lack of setae on the openings of the penis lobes.The nymph is easily distinguished from Tepakia by its narrow gills and from New Zealand genera with narrow gills as follow: from Atalophlebioides Phillips, Neozephlebia Penniket, Austroclima Towns & Peters and Mauiulus Towns & Peters by the shallow anteromedian emargination of the labrum with subequal-sized denticles; and from Cryophlebia Towns & Peters and Austronella Towns & Peters by the presence of posterolateral projections on abdominal segments 2, 3, 4 or 5-9.