Zootaxa 3948 (1): 109–124
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Copyright © 2015 Magnolia Press
Article
ISSN 1175-5326 (print edition)
ZOOTAXA
ISSN 1175-5334 (online edition)
http://dx.doi.org/10.11646/zootaxa.3948.1.7
http://zoobank.org/urn:lsid:zoobank.org:pub:60224FB0-4E2D-4892-89E2-9F3160180C91
Description of immatures associated with adults of two species of Macrogynoplax
Enderlein and one species of Enderleina Jewett (Plecoptera: Perlidae)
of the Brazilian Amazon
JOSÉ MOACIR FERREIRA RIBEIRO1,2 & INOCÊNCIO DE SOUSA GORAYEB2
1
Laboratório de Biologia Aquática, Departamento de Ciências Biológicas, Universidade Estadual Paulista “Júlio de Mesquita Filho
(UNESP), Assis, São Paulo, Brazil. E-mail: [email protected] / [email protected]
2
Museu Paraense Emílio Goeldi, Coordenação de Zoologia, Caixa Postal 399, Belém, Pará, Brasil. E-mail: [email protected]
Abstract
Nymphs of Macrogynoplax delicata Ribeiro-Ferreira & Froehlich, 1999, M. pulchra Ribeiro-Ferreira & Froehlich, 1999,
and E. froehlichi Ribeiro-Ferreira, 1996 were associated with adults by rearing and described for the first time. The known
distribution of each species has been increased.
Key words. Stonefly, immature, rearing, aquatic insect, South America
Introduction
The perlid genus Macrogynoplax Enderlein, 1909 is known from Suriname, Guyana, Venezuela, Peru, and Brazil.
The 14 known species occur at both high and low altitudes. However, the genus is apparently uncommon in the
Central Amazon (Ribeiro-Ferreira & Froehlich 1999, Bobot & Hamada 2002, Ribeiro & Rafael 2007, Fochetti &
Tierno de Figueroa 2008, Froehlich 2010, 2012). Seven species of Macrogynoplax have been previously recorded
from Brazil and six of these occur in the Brazilian Amazon (Stark et al. 2009, Froehlich 2010).
For most species, as in other perlid genera, species descriptions are based only on adults without immature
stages associated. Only a few publications include descriptions of nymphs. The lack of keys for identification of
nymphs of Macrogynoplax has hindered the study of the systematics and ecology of this genus in the Brazilian
Amazon (Froehlich 1984 a, b, Olifiers et al. 2004). Only two studies have described nymphs of Macrogynoplax
from Brazil, Froehlich (1984a) (M. veneranda Froehlich) and Stark & Zwick (1989) (M. neblina Stark). In the
present study, nymphs of two species of Macrogynoplax are described and associated with adults for the first time.
This increases the number of known nymphs of Macrogynoplax from the Brazilian Amazon to four. Previous
studies in which Macrogynoplax species were described include Ribeiro & Rafael (2007) (M. anae), RibeiroFerreira & Froehlich (1999) (M. delicata), Stark (1996) (M. flinti), Zwick in Stark & Zwick (1989) (M. geijskesii),
Enderlein (1909) (M. guayanensis), Stark (1996) (M. kanuku), Bispo & Neves in Bispo et al. (2005) (M.
matogrossensis), Stark in Stark & Zwick (1989) (M. neblina), Ribeiro-Ferreira & Froehlich (1999) (M. poranga),
Ribeiro-Ferreira & Froehlich (1999) (M. pulchra), Stark in Stark & Zwick (1989) (M. spangleri), Stark 1996 (M.
truncata), Froehlich (1984b) (M. veneranda), and Stark, (1996) (M. yupanqui).
The genus Enderleina Jewett (1960) is known from Guyana, Venezuela, Peru, and northern Brazil (states of
Roraima and Amazonas). Of the six Neotropical species known, four occur in the Brazilian Amazon. Studies in
which species of Enderleina were described include Stark (1989) (E. bonita), Stark (1989) (E. flinti), RibeiroFerreira (1996) (E. froehlichi), Jewett (1960) (E. preclara), Stark (1989) (E. yano), and Derka & Tierno de
Figueroa (2013) (E. khazeni). With this study the number of Enderleina nymphs known from the Brazilian Amazon
is increased to two.
Accepted by B. Kondratieff: 26 Mar. 2015; published: 20 Apr. 2015
109
Material and methods
The material examined this study was borrowed from the Instituto Nacional de Pesquisas da Amazônia (INPA),
Manaus, Amazonas, Brazil and Museu Paraense Emílio Goeldi (MPEG), Belém, Pará, Brazil. Additional material
was collected from several states of the Brazilian Amazon at the following locations: Floresta Nacional do Amapá
(Programa de Pesquisa em Biodiversidade PPBio: Núcleo Leste do Pará and Amapá), Reserva Florestal Adolpho
Ducke (Amazonas), Peixe-Boi River Basin and São Geraldo do Araguaia, Serra dos Martírios-Andorinhas.
Nymphs were either preserved in 70% to 96% alcohol; others were reared directly in the streams or in tanks in the
laboratory. Descriptions and diagnosis follow the terminology used by Shepard & Stewart (1983), Olifiers et al.
(2004), Sandberg (2011), and Stewart & Stark (2002).
Result and discussion
Macrogynoplax delicata Ribeiro-Ferreira & Froehlich, 1999
(Figs. 1–11)
Macrogynoplax delicata Ribeiro-Ferreira & Froehlich 1999: 134, Figs. 1–5; Ribeiro & Rafael 2007: 19, Figs. 1, 4–5, 9–17;
Froehlich 2003: 133 (key); Ribeiro et al. 2009: 34 (catalogue); Froehlich 2010: 184 (catalogue).
Macrogynoplax marauia Froehlich, 2003: 133, p. 132, figs. 15–18. synonym of M. delicata, Ribeiro & Rafael 2007: 19.
Diagnosis of nymph. Body light brown. Frons light brown, labrum light yellow; parietália yellowish, light brown
at epicranial line. Pronotum light brown, yellowish laterally.
Description of nymph. Female: Body robust, length 9.0–10.0 mm (not including cerci, n=20), general color
shiny dark brown to slightly yellowish; body robust (Fig. 1). Head: Antennae dark yellow, basally with dark
margin, scape as wide as long. Occiput in dorsal view without spots; frons dark brown, M-line distinct, with small
yellow spots lateral to ocelli. Parietalia dark yellow, brown at coronal line and post-frontal line. Eyes black;
distance between ocelli 0.4 mm, from ocelli to inner margin of eye, 0.6 mm (Fig. 2). Mouthparts whitish yellow.
Labrum light yellow ventrally, and dark yellow on dorsum with thin setae (Fig. 3). Maxillae greenish-yellow, cardo
and stipes sclerotized, light yellow, medial margin of stipes sclerotized, dark yellow, and with groups of short
subequal robust setae; lacinia with two light brown teeth and a row of six robust, long bristles below teeth (Fig. 4).
Mandible whitish yellow with five acute teeth on distal margin, decreasing in size to base, but 4th tooth larger than
the 3rd, 5th and 2nd and 4th tooth subequal; two rows of bristles on the ventral margin; first row starting at the base of
the 2nd tooth and other at base of the 4th tooth (Figs. 5, 6). Pronotum dark brown, without median spots, posterior
margin with median concavity, pronotal sulcus narrow and band lateral wide; ventral side whitish yellow, with
sulcus well-marked; angle between the anterior arms of Y-line narrow, without groups of setae on the anterior side
or lateral of the Y-line (Figs. 2, 8). Meso- and metanotum light brown, mesonotum with median light yellow area,
with yellowish bands laterally, covered with thin dark pilosity; lateral border, light yellow, spots narrow and long,
dark brown pigment extends laterally from apex to base; metanotum similar to mesonotum with variably sized
medial spots (Fig. 7). Gills present on ASC [1, 2, 3], PSC [1, 2] with simple branches; PSC [3] with double
branches; AT [2, 3] with triple branches and SL with multiple branches (Fig. 8). Prothoracic leg raptorial, light
yellow, unpigmented ventrally; coxa with few bristles; trochanter with stout bristles stronger and dense in posterior
region; femur light yellow without transverse bristles apically; with bristle short and strong and a fringe of bristle
on the dorsoanterior margin; ventral margin with long and sparse bristles and a lobular projection at the distal third,
short in dorsal view; tibiae without short band of bristles on anterior and posterior margins; a cluster of fine and
dense whitish bristles along of entire segment, posterior margin with three large, strong spines distally; tarsus pale
yellow dorsally and ventrally, with bristles fine and long (Figs. 9, 10). Abdomen dorsally light brown, similar to
head and thorax; intersegmental membranes darker; segments with complete dorsal row of bristles that at posterior
margin becoming more distinct on sterna 8–10. Ventrally, bristles extend to a median gap equal to of sternum width
on segments 1–4; bristle row complete on sterna 8–10, eight sternite, with a median U-shaped, short notch; sterna
with sparse bristles, posterior three sterna with long bristles distally. Cerci dark to light yellow (Fig. 11).
Male. Unknown.
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RIBEIRO & GORAYEB
FIGURES 1–6. Female Macrogynoplax delicata. 1—body; 2—head and pronotum, dorsal view; 3—head, ventral view; 4—
maxillae and lacinia; 5—mandible, dorsal view; 6—mandible, ventral view (photos: J.M.F. Ribeiro).
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FIGURES 7–11. Female Macrogynoplax delicata. 7—wing pad; 8—sternal sulci and gills; 9–10—legs, dorsal and ventral
views; 11—Abdomen, ventral view. (photos: J.M.F. Ribeiro).
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RIBEIRO & GORAYEB
Variation: Female. Body color ranging from light brown to light yellow with a greenish cast; head with line
epicranial well defined and distinct; head without spots and pronotum, meso- and metanotum with light yellow
spots; median area with anterior and posterior brown bands; lateral spots variable in length; sterna whitish yellow.
Long bristles present on lateral margins of terga 5–10 and on median-lateral margins of sterna 6–10.
Distribution. BRAZIL, Amazonas, Manaus Municipality, Reserva Florestal Adolpho Ducke; Amapá,
Macapá Municpality, Floresta Nacional do Amapá (new record); Pará, Peixe-Boi Municipality, Peixe-Boi River
Basin, Nova Timboteua Municipality (new record) and São Geraldo do Araguaia Municipality, Serra dos
Martírios-Andorinhas (new record).
Remarks. The association with the adult was confirmed by rearing. The nymph differs from M. pulchra by the
shiny dark brown color and more robust habitus, the head dark brown, M-line conspicuous, the presence of a slight
median concavity on the pronotum, the angle of the anterior arms of the Y-line of mesosternum long, femur
uniformly dark yellow to dark brown with a short lobular projection in dorsal and ventral view, and the welldefined dark spots on the wing pads.
Macrogynoplax delicata was the frequently collected species during this study from two river basins of the
Reserva Florestal Adolpho Ducke Manaus, Amazonas, two streams in the Floresta Nacional do Amapá, two
streams of the Peixe-Boi River Basin, and a stream of Serra dos Martírios-Andorinhas, northeastern and southern
of state of Pará. We identified additional adults from the following localities, Reserva Florestal Adolpho Ducke
(Manaus, Amazonas, several adults, males and females) INPA and Fazenda Candiru, adult male MPEG.
Material examined. BRASIL, Amazonas State Reserva Florestal Adolpho Ducke (RFAD), Am-010, Km 30,
Barro Branco stream, [aquatic net], 14.vi.1989, L.C. Machado & R.S. Rocha (3 nymphs, INPA); same location,
Centro de Instrução de Guerra na Selva (SIGS), Km 28, Branquinho stream, [aquatic net], 16.vi.1993, A.C.
[Ribeiro-Ferreira] (3 nymphs, INPA); same location, [aquatic net], 16.vi.1993, A. Celeste (3 nymphs, INPA); same
location, (RFAD), Barro Branco stream, [aquatic net], 1995, [A. Celeste] (2 nymphs, INPA); same location,
[aquatic net], 02.vii.1995, A. Celeste (13 nymphs, INPA); same location, [aquatic net], 11.ix.1995, [A. Celeste] (1
nymphs, INPA); same location, [aquatic net], v. 1995, [A. Celeste] (10 nymphs, INPA); same location, Bons
Amigos stream, [aquatic net], 24.iii. 1995, (reared in laboratory), A. Celeste (1 exuvia, 1 adult, INPA); same
location, [aquatic net], 06.vi.1995, A. Celeste (1 exúvia, INPA); same location, [aquatic net], 24.iv.1996, S. Cleto
(8 nymphs, INPA); same location, Am-010, Km 26, aquatic net, 01.iii.2000, E.C. Litaiff (1 nymph, INPA); same
location, Projeto Dinâmica Biológica de Fragmentos Florestais (PDBFF), Fazenda Dimona, (Rio Cuieiras), leaf
litter [aquatic net], 06.ii.2001, J.L. Nessimian (3 nymphs, INPA); same location, Fazenda Dimona, (Rio Cuieiras),
leaf litter in riffles, [aquatic net], 07.ii.2001, J.L. Nessimian (1 nymph, INPA); same location, PDBFF, Fazenda
Porto Alegre, (Rio Urubu), leaf litter in riffles, [aquatic net], 08.ii.2001, J.L. Nessimian (1 nymph, INPA); same
location, Fazenda Esteio, leaf litter in riffles, [aquatic net],10.ii.2001, J.L. Nessimian (1 nymph, INPA); same
location, (Rio Urubu), [aquatic net], 11.ii.2001, J.L. Nessimian (1 nymph, INPA); same location, in riffles [aquatic
net], 11.ii.2001, J.L. Nessimian (1 nymph, INPA); same location, Km-41, [aquatic net], 11.ii.2001, J.L. Nessimian
(1 nymph, INPA); same location, ZF3, Km 23, Fazenda Esteio/Colosso, (Rio Urubu), 12. ii. 2001, [aquatic net],
J.L. Nessimian (3 nymphs, INPA); same location, (Rio Preto da Eva), leaf litter in riffles, 12. ii. 2001, [aquatic net],
J.L. Nessimian (1 nymph, INPA); same location, Fazenda Esteio/Gavião, J.L. Nessimian (2 nymphs, INPA); same
location, [aquatic net], 12. ii. 2001, J.L. Nessimian (1 nymph, INPA); same location, Reserva [Florestal Adolpho]
Ducke, Acar stream, 13.x. 2001, [aquatic net], Ribeiro, J.M.F. (2 nymphs, INPA); same location, PDBFF, Fazenda
Esteio/Gavião, (Rio Urubu), leaf litter in riffles, 20.x.2001, [aquatic net], J.L. Nessimian (10 nymphs, INPA); same
location, Fazenda Esteio/Dimona, (Rio Cuieiras), 21.x.2001, [aquatic net], J.L. Nessimian (2 nymphs, INPA); same
location, 22.x.2001, in riffles, [aquatic net], J.L. Nessimian (1 nymph, INPA); same location, Fazenda Esteio/C.
Powell, 23.x.2001, [aquatic net], J.L. Nessimian (2 nymphs, INPA); same location, Reserva do Gavião, 25.x.2001,
[aquatic net], J.L. Nessimian (1 nymph, INPA); same location, Br-174, km-7, Fazenda Dimona, in riffles,
07.ii.2002, J.L. [aquatic net], Nessimian (1 nymph, INPA); same location, Reserva [Florestal Adolpho] Ducke,
Ipiranga stream, [aquatic net], 13.ix.2002, J.M.F. Ribeiro & J. Vidal (1 nymph, INPA); same location, stream of
ounce, [aquatic net], 21.v. 2002, Ribeiro, J.M.F. Ribeiro & F.P. Gouveia (1 nymph, INPA); same location, Reserva
[Florestal Adolpho] Ducke, Ipiranga stream, [aquatic net], 12.x.2002, J.M.F. Ribeiro (4 nymphs, INPA); same
location, [aquatic net],15.ix.2002, J.M.F. Ribeiro & J. Vidal (4 nymphs, INPA); same location, Reserva de
Desenvolvimento Sustentável do Tupé stream of riffles, leaf litter [aquatic net], 17.viii.2002, (1 nymph, INPA);
same location, ZF3, Fazenda Esteio, Ponta Verde stream, 12.vi.2006, leaf litter of S. bifida, J.D. Paula (1 nymph,
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INPA); same location, Reserva [Florestal Adolpho] Ducke, Ipiranga stream, [aquatic net], 19.iv.2003, Ribeiro,
J.M.F (1 exuvia, INPA); same location, PDBFP, [aquatic net], 18–24.ii.2003 (3 nymphs, INPA); same location,
Universidade Federal do Amazonas (UFAM) stream (reared), 12.xi.2007, F.M. Silva & J.M.F. Ribeiro (1 nymph in
transition of adult, INPA); same location, UFAM stream (reared), 26.v.2007, F.M. Silva (1 nymph, 1 adult, INPA);
same location, Barro Branco stream, [aquatic net], 31.xi.2007, F.M. Silva (1 nymph, INPA); same location, UFAM
stream (reared), 21.v.2008, F.M. Silva (3 nymphs, INPA); same location, [aquatic net], 26.v.2008, F.M. Silva (1
nymph, INPA); same location (reared), [aquatic net], 05.ix.2008, F.M. Silva (1 nymph, INPA); same location,
Acará stream, [aquatic net], 17.viii.2011, J.M.F. Ribeiro (1 nymph, INPA); Pará State, Nova Timboteua
Municipality, Burrinho stream, 12.iii.2010, J.M.F. Ribeiro & M.J. Candido (3 nymphs, MPEG); same location,
Peixe-Boi, Municipality, sieve, dragnet, 17.iii.2011, 2 BIS stream, J.M.F. Ribeiro & Monteiro-Junior (3 nymphs,
MPEG); same location, São Geraldo do Araguaia, Municipality, Serra dos Martírios-Andorinhas, 30.x.2011,
[aquatic net, sieve, dragnet], Água Verde stream, 06 0539,6´´S 48 2919,9´´W, J.M.F Ribeiro & C.R.M. Santos (1
nymph, MPEG). Amapá, Macapá, Municipality, Floresta Nacional do Amapá, PPBio, Japiim stream, between the
trails I e II, (00o58’37,5’’S 51o36’50 W), [aquatic net, sieve, dragnet], 22–30.x.2009, J.M.F. Ribeiro & C.R.M.
Santos (5 nymphs, MPEG).
Ecological notes. Nymphs of M. delicata were collected from rocky substrates with leaf packs, most
commonly associated with leaf litter, but sometimes found on submerged logs in riffle reaches. Nymphs could be
collected from substrates as described above from pristine streams with little sedimentation to those that have been
impacted by deforestation. Adults of this species were rarely collected even when using light traps. Physicochemical parameters of selected streams are presented in Table 1.
TABLE 1. Physicochemical parameters of streams in Brazil where Macrogynoplax delicata was collected.
Parameters
BB
2BIS
AV
JA
Water temperature ( C)
26.0
26.0
24.0
24.0
pH
4.5
6.0
6.7
5.4
Specific conductance (uS/cm)
13.5
12.0
15.4
13.6
Dissolved oxygen (mg/l)
46.0
6.8
27.0
2.8
o
Width (m)
3.0
2.5
2.7
3.5
Depth (cm)
0.75
0.36
0.24
0.4
0.6
0.25
0.31
0.6
1.35
0.25
0.31
0.9
Velocity (m/s)
3
Discharge (m /s)
BB = Amazonas, Manaus, Reserva Florestal Adolpho Ducke, Barro Branco Stream; 2BIS = Pará, Peixe-Boi River basin,
second BIS Stream; AV = Pará, Serra dos Martírios-Andorinhas, Água Verde Stream, JA = Amapá, Floresta Nacional do
Amapá, Japiim Stream.
Macrogynoplax pulchra Ribeiro-Ferreira & Froehlich, 1999
(Figs. 12–22)
Macrogynoplax pulchra Ribeiro-Ferreira & Froehlich, 1999:136, figs. 7–10; Ribeiro & Rafael, 2007:23, Figs. 2, 3, 7, 26–34;
Ribeiro et al., 2009:35 (catalogue); Froehlich, 2010 (catalogue).
Diagnosis of nymph. Body dark yellow, slightly greenish. Frons dark brown and labrum, light brown; parietália
dark yellow, light brown at epicranial line. Pronotum light brown, yellowish laterally. Mesonotum and metanotum
brown without yellowish muscle scars; lateral spots not elongated.
Description of nymph. Female: Body length 9.0 mm (not including the cerci, n=10), general color dark
yellow (Fig. 12). Antennae dark yellow, basally with yellowish margin; scape longer then wide. Occiput in dorsal
view without spots. Frons dark brown to dark yellow; M-line distinct, with small yellow spots lateral to ocelli.
Parietalia dark yellow, more distinct at coronal line and post-frontal line. Eyes black; distance between ocelli 0.4
mm, from ocelli to inner margin of eye, 0.7 mm (Fig. 13). Mouthparts dark yellow. Labrum ventrally light yellow,
dorsum light brown with small bristles at edge (Fig. 14). Maxillae light brown; sclerotized outer part, cardo and
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stipes, light brown, medial margin of stipes sclerotized the same color, with groups of long, robust setae; lacinia
pale with two light brown teeth; a row of, nine robust long bristles below inferior tooth (Figs. 15). Mandible, pale
with five acute teeth on distal margin; teeth decreasing in size to base, but the 4th tooth larger than the 2nd and 5th, 2nd
and 4th teeth subequal; two rows of long bristles on ventral mandibular margin, first row starting at base of 2nd tooth
and 2nd row beginning at base of 4th tooth (Figs. 16, 17). Pronotum dark brown, without medial spots; posterior
margin with median concavity; pronotal sulcus narrow, lateral band wide and posterior margin yellowish, ventrally
dark yellow with sulcus well-marked; angle between the anterior arms of Y-line narrow, without groups of setae
anteriorly and laterally of Y-line (Figs. 13, 19). Meso-and metanotum dark yellow to light brown; mesonotum with
median area dark yellow, laterally with whitish yellow bands, covered laterally with thin dark pilosity; lateral
border light yellow without spots; Metanotum similar to mesonotum but with fewer lateral bristles (Fig. 18). Gills
present on ASC [1, 2, 3], PSC [1, 2] with simple branches; PSC [3], with double branches; AT [2, 3], with branches
triple and SL with multiple branches (Fig. 19). Prothoracic leg raptorial, dark yellow, unpigmented ventrally; coxa
with few bristles, trochanter with bristles stronger and dense posteriorly; femur dark yellow, without transverse
bristles but with long anterodorsal fringe of whitish bristles and a lobular projection at the distal ⅓; projection long
in dorsal view. Tibiae without band of short bristles on anterior and posterior margin but with a long fringe of fine
and dense whitish bristles along entire segment, posterior margin with three large, strong spines distally; tarsus pale
yellow, with fine and long bristles (Figs. 20, 21). Abdomen dark yellow dorsally, intersegmental membranes darker
brown; segments with complete dorsal row of bristles at posterior margin, more distinct on sterna 8–10; ventrally,
bristles extend to a median gap equal to of sternum width on the segments 1–4, 1/5 of sternum 5–6, and bristle row
complete on sterna 7–10; sterna 8–10 with dense, short bristles; sternite 8 with a median short U-shaped notch.
Cerci dark brown to light brown (Fig. 22).
Male: Male nymph differs from female nymph by lack of the median U-shaped notch on sternite 8. Body
length 7.5 mm, head 0.9 mm long and 2.0 mm wide. Distance between ocelli 0.4 mm; from each ocelli to inner
margin of eye 0.7 mm. Pronotum light yellow to yellow-orange, 1.1 mm long, 2.6 mm wide; mesonotum (wing pad
I) 1.2 mm long, 2.1 mm wide; metanotum (wing pad II) 1.2 mm long, 2.0 mm wide.
Variation. Females: Body color ranging from dark yellow to light brown (with greenish cast). Pronotum
lacking spots. Dense short bristles sometimes present on lateral margins of terga 3–10 and along median-lateral
margins of sterna 2–10. In some immature nymphs M-line and epicranial suture on posterior portion the head
evident. Immature nymphs have denser pilosity.
Male. Body more shiny and robust. Color varies from light yellow in smaller nymphs to yellow orange in
mature nymphs. Without spots in posterior portion of the head. Pro-, meso-, and metanotum with spots evident
(muscle scars) but these are less conspicuous in immature nymphs. More bristles occur on smaller nymphs, mainly
on the pronotum, mesonotum, and legs.
Distribution. BRAZIL, Amazonas, Manaus (Reserva Florestal Adolpho Ducke). Amapá State (Floresta
Nacional do Amapá), (new record) and Pará State (Capanema), (new record).
Remarks. The association with the adult was confirmed by rearing. The nymph differs from M. delicata by the
more robust body slightly more shiny dark yellow to yellow-greenish (light yellow to orange in male nymphs)
coloration. Head dark brown in front and dark yellow medially; M-line conspicuous. A deep medial concavity
occurs on the pronotum. The angle of the anterior arms of the Y-line of mesosternum is longer. The femur is
uniformly dark yellow to light brown and the long lobular projection in dorsal and ventral view has a pattern of
dense bristles and a long fringe. Spots on the wing pad uniform, without laterally elongated dark brown spots.
Macrogynoplax pulchra is a species that has been collected not as frequently as M. delicata. Specimens were
collected from two river basins in the Reserva Florestal Adolpho Ducke, Manaus, Amazonas State and from a
tributary of the Rio Negro, Amazonas (by E. Fittkau), two streams in the Floresta Nacional do Amapá, two streams
of the Peixe-Boi River Basin, and Capanema in northeastern Pará State. We identified additional male and female
adults from localities that included Reserva Florestal Adolpho Ducke, Manaus, Amazonas State (INPA) and four
male adults and four female adults (MPEG).
Material examined. BRASIL, Amazonas State, 23.ii.1987, W.L.S. Costa (1 nymph, INPA); same location,
Reserva [Florestal Adolpho] Ducke (RFAD), Am-010, km-30, 14.vi.1989, Barro Branco stream, [aquatic net], L.C.
Machado & R.S. Rocha (1 nymph, INPA); same location, Bons Amigos stream, [aquatic net], vii.1994, A. Celeste,
(7 nymphs, INPA); same location, Barro Branco stream, [aquatic net], 02.vii.1995, A.C. Celeste (1 nymph, INPA);
same location, (reared), 11.vii.1995, A.C. Ferreira (1 exuvia, 1 adult, INPA); same location, (reared), 24.iv.1995,
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FIGURES 12–17. Female Macrogynoplax pulchra. 12—body; 13—head and pronotum, dorsal view; 14—head, ventral view;
15—maxillae and lacinia; 16—mandible, dorsal view; 17—mandible, ventral view (photos: J.M.F. Ribeiro).
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FIGURES 18–22. Female Macrogynoplax pulchra. 18—wing pad; 19—sternal sulci and gills; 20–21—legs, dorsal and ventral
views; 22—Abdomen, ventral view. (photos: J.M.F. Ribeiro).
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A.C. Ferreira (1 exuvia, 1 adult, INPA); same location, Projeto de Dinâmica Biológica de Fragmentos Florestais
(PDBFF), ZF3, Km-23, Fazenda Esteio (022355S-595242W), leaf litter [aquatic net], 23.x.2001, J.L. Nessimian (2
nymphs, INPA); same location, (River Urubu), J.L. Nessimian (1 nymph, INPA); same location, in riffles [aquatic
net], J.L. Nessimian (1 nymph, INPA); same location, in riffles, 11.ii.2001, [aquatic net], J.L. Nessimian (1 nymph,
INPA); same location, 21.x.2001, in riffles, [aquatic net], J.L. Nessimian (1 nymph, INPA); same location, (RFAD),
Ipiranga stream, [aquatic net], 15.ix.2002, J.M.F Ribeiro & J. Vidal (1 nymph, INPA); same location, 12.x.2002,
J.M.F. Ribeiro, (1 nymph, INPA); same location, Tinga stream, aquatic net, 08–11.xi.2002, J.M.F. Ribeiro (1
nymph, INPA); same location, aquatic net, 28.vi.2003, J.M.F. Ribeiro, (1 nymph, INPA); same location, River
Negro, E. Fittkau, (1 nymph, INPA); same location, E. Fittkau (1 nymph, INPA); Amapá State, Floresta Nacional
do Amapá, plote-PPBio, Japiim stream, between the trails I e II, (00o58’37,5’’S 51o36’50 W), [sieve and trawl], 22–
30.x.2009, Ribeiro, J.M.F. & C.R.M. Santos (2 nymphs, MPEG). Pará State, Capanema municipality, Piranguinha
streams, in riffles, [aquatic net], 14.xii.2011, J.M.F. Ribeiro, D.A. Silva & M.L.S. Silva (3 nymphs, MPEG); same
location, leaf litter, 22.viii.2011, aquatic net, J.M.F. Ribeiro, D.A. Silva & M.L.S. Silva (3 nymphs, MPEG).
Ecological notes. Nymphs of M. pulchra were usually associated with leaf litter in riffles with gravel substrate
mixed with small stones. Nymphs have also been collected from submerged wood along riffle areas. This species
apparently is restricted to relatively pristine streams in areas that have not been impacted by anthropogenic
disturbances. However, stream in areas of subsistence agriculture still support this species. Physico-chemical
parameters associated with M. pulchra are given in Table 2.
TABLE 2. Physicochemical parameters of streams in Brazil where Macrogynoplax pulchra was collected.
Parameters
BB
JA
PI
Water temperature ( C)
24.8
24.0
25.9
pH
6.1
5.4
5.8
Specific conductance (uS/cm)
13.5
13.6
124.2
Dissolved oxygen (mg/l)
46.0
2.8
4.7
Width (m)
3.0
3,5
2.0
Depth (cm)
0.60
0.4
0.4
o
Velocity (m/s)
1.15
0.6
0.6
3
2.07
0.9
0.4
Discharge (m /s)
BB = Amazonas, Manaus, Reserva Florestal Adolpho Ducke, Barro Branco Stream; JA = Amapá, Floresta Nacional do
Amapá, Japiim Stream; PI =Pará, Capanema, Piranguinha stream.
Enderleina froehlichi Ribeiro-Ferreira, 1996
(Figs. 23–33)
Enderleina froehlichi Ribeiro-Ferreira, 1996:145, figs. 1–6; Ribeiro & Rafael, 2005:62, Figs. 1–5, 8–15; Ribeiro et al.,
2009:34; Stark et al., 2009:120; Froehlich, 2010:178 (catalogue).
Diagnosis of nymph. Body dark yellow. Frons dark yellow and labrum light yellow with anterior margins light.
Parietália dark yellow more pronounced at epicranial line. Pronotum light yellow medially, whitish yellow
laterally. Meso-and metanotum dark yellow with faint muscle scars but without laterally elongated spots.
Description of nymph. Female: Body length 6.0 mm–11.0 mm (not including cerci, n=10). Body robust and
shiny light yellow (Fig. 23). Antennae light yellow, basally with darker margin; scape longer then wide. Occiput
dorsally without spots. Frons light yellow to dark yellow; M-line not distinct, without small yellow spots laterally
to ocelli. Parietalia dark yellow, more distinct at coronal line and post-frontal line. Eyes black; distance between
ocelli 0.3 mm, from ocelli to inner margin of eye, 0.6 mm (Fig. 24). Mouthparts dark yellow. Labrum light yellow
ventrally, dorsum dark yellow with small bristles forming fringe (Fig. 25). Maxillae dark yellow; sclerotized outer
part, cardo and stipes dark yellow; medial margin of stipes sclerotized the same color with groups of long, thin
setae; lacinia dark yellow, with two dark brown teeth and a row of five robust long bristles below inferior tooth and
eight short bristles near the base (Fig. 26). Mandible light brown with five acute teeth on distal margin, decreasing
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in size to base, but 4th tooth larger than the 2nd, 3rd and 5th, 2nd and 4th tooth subequal. Two rows of bristles on the
ventral margin forming fringe from first row starting at the base of 2nd tooth, 2nd row beginning at base of the 4th
tooth with long bristles; a group of fine long bristles occurs on the anterolateral margin of the dorsal and ventral
surfaces of the region (Figs. 27, 28). Pronotum dark yellow, without median spots; posterior margin with median
concavity; pronotal sulcus narrow and lateral band wide; ventrally dark yellow with sulcus well marked; angle
between the anterior arms of Y-line narrow and without groups of setae on anterior edge and lateral of the Y-line
(Figs. 24, 30). Meso- and metanotum dark yellow; mesonotum with median area yellowish, with milky white
yellow bands lateral borders; mesonotum covered with thin brown pilosity, lateral border yellowish without spots;
metanotum similar to mesonotum but, with more bristles laterally (Fig. 29, 30). Gills present on ASC and PSC [1,
2, 3], AT [2, 3], PT [3] all of with simple branches and SL with multiple branches (Fig. 30). Prothoracic leg light
yellow dorsally, unpigmented ventrally; coxa with bristles long and robust; trochanter with bristles stronger and
dense; femur with a long fringe of distal whitish bristles along anterodorsal margin; ventral margin with bristles
long and robust distally and without lobular distal projection; ventral sulcus extending from apex to near base.
Tibiae with band of bristles short and robust on the posterior margin and with a long fringe of fine and dense
whitish bristles along entire segment, posterior margin with four large, strong spines distally. Tarsus pale yellow
with fine, long bristles (Figs. 31, 32). Abdomen dark yellow dorsally, intersegmental membranes darker brown;
segments with complete dorsal row of bristles at posterior margin, more distinct on sterna 3–10. Ventrally, bristles
extend to median gap equal to of sternum width on the segments 1–3, 1/5 of sternum 4–5, and bristle row complete
on sterna 6–10. Sterna with long and short bristles; sternite 8, with a shallow median U-shaped notch. Cerci dark
brown to light brown (Fig. 33).
♂. Unknown.
Variations. Not observed.
Distribution. BRAZIL, Amazonas, Manaus (Reserva Florestal Adolpho Ducke, Universidade Federal do
Amazonas, stream, Reserva de Desenvolvimento Sustentável do Tup) Rio Negro and Amapá (Serra Lombard,
Limão).
Remarks. The nymphal-adult association was confirmed by rearing. The nymph differs from E. flinti by
general color with the body being light yellow and the head dark yellow. The M-line inconspicuous, the lacinia has
a slightly rounded base; the presence of a large and deep median concavity occurs on the pronotum. The angle of
the anterior arms of the Y-line of mesosternum is longer; femora are uniformly dark yellow with a ventral sulcus
extending from apex to near the base and with short and dense bristles and a long fringe in dorsal view. Spots are
absent on the wing pads.
Four species of Enderleina are recorded for northern Brazil and Brazilian Amazon (E. flinti, E. yano, E.
preclara and E. froehlichi). However, only the nymph of E. flinti has been provisionally described by Stark (1989)
collected in the Serra da Neblina. This association was based solely on distribution, without nymph-adult
association.
Enderleina froehlichi is considered a rare species with a restricted distribution to the Brazilian Amazon. Two
nymphs collected by E. Fittkau with the locality of Rio Negro, Amazonas were identified from the collection of
Aquatic Insects of Museu de Zoologia da Universidade de São Paulo (MZUSP) through courtesy of the curator,
Dra. Sônia Casari. One adult collected by J. Bechyné from Serra Lombard, Limão, Amapá was identified from the
collection of Museu Paraense Emílio Goeldi. The remainder of the material examined was borrowed from INPA
collected in Reserva Florestal Adolpho Ducke and in the vicinity of the city of Manaus.
Material examined. BRASIL, Amazonas, Manaus, Universidade Federal do Amazonas (UFAM), (reared),
09.ix.2008, J.M.F. Ribeiro & F.M. Silva (1 nymph, INPA); same location, Reserva [Florestal Adolpho] Ducke
(RFAD), Barro Branco stream, [aquatic net], 18.vii.2003, J.M.F. Ribeiro (1 nymph, INPA); same location, Parque
Municipal do Mindú stream I, 05.x.1993 (1 nymph, INPA); same location, Reserva de Desenvolvimento
Sustentável do Tupé (RDST), in leaf litter, 14.x.2002, (4 nymphs, INPA); same location, in riffles with leaf litter,
18.v.2002 (6 nymphs, INPA); same location, (RFAD), Barro Branco stream, (reared), 18.vii.2008, J.M.F. Ribeiro (1
exuvia, INPA); same location, Projeto de Dinâmica Biológica de Fragmentos Florestais (PDBFF), ZF3 Km-7,
Fazenda Porto Alegre, stream in river Urub, (2o21’11’’S–59o 19’1’’W), 08.ii.2001, J.L. Nessimian (1 nymph,
INPA); same location, stream in UFAM, rearing in tank, 10.xi.2008, J.M.F. Ribeiro & F.M. Silva (1 nymph, INPA);
same location, rearing in tank, 05.ii.2008, J.M.F. Ribeiro & F.M. Silva (1 nymph, 1, adult, 1 exuvia, INPA); same
NYMPHS OF MACROGYNOPLAX AND ENDERLEINA
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FIGURES 23–28. Female Enderleina froehlichi. 23—body; 24—head and pronotum, dorsal view; 25—head, ventral view;
26—maxillae and lacinia; 27—mandible, dorsal view; 28—mandible, ventral view (photos: J.M.F. Ribeiro).
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FIGURES 29–33. Female Enderleina froehlichi. 29—wing pad; 30—sternal sulci and gills; 31–32—legs, dorsal and ventral
views; 33—Abdomen, ventral view. (photos: J.M.F. Ribeiro).
NYMPHS OF MACROGYNOPLAX AND ENDERLEINA
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location, stream in UFAM, 20.vi.2008, J.M.F. Ribeiro & F.M. Silva (1 nymph, INPA); same location, (RFAD),
Barro Branco stream, 15.vii.2008, rearing in tank, J.M.F. Ribeiro & F.M. Silva (1 nymph, 1 adult, 1 exuvia, INPA).
[Brasil, Amazonas], Rio Negro, E. Fittkau, (2 nymph, MZUSP). Amapá State, Serra Lombard, Limão, J. Bechyné,
03.ix.1961 (1 female, adult, MPEG).
Ecological notes. Nymphs of E. froehlichi were collected from sandy and rocky substrates and leaf litter from
relatively pristine streams with intact riparian zones. Adult are rarely observed in the field, but can be collected by
using light traps. Physico-chemical parameters associated with E. froehlichi are presented in Table 3.
TABLE 3. Physicochemical parameters of streams in Brazil where Enderleina froehlichi was collected.
Parameters
BB
o
Water temperature ( C)
24.8
pH
6.1
Specific conductance (uS/cm)
13.5
Dissolved oxygen (mg/l)
46.0
Width (m)
3.0
Depth (cm)
0.60
Velocity (m/s)
1.15
3
2.07
Discharge (m /s)
BB = Amazonas, Manaus, Reserva Florestal Adolpho Ducke, Barro Branco Stream.
Comments about the genus Macrogynoplax and Enderleina in the Brazilian Amazon
The South America Plecoptera fauna includes representatives of six families, Austroperlidae, Eustheniidae,
Diamphipnoidae, Gripopterygidae (Antarctoperlaria); Notonemouridae, and Perlidae (Arctoperlaria). All genera
within these families are endemic to South America with the exception of the perlid genus Anacroneuria, which
also occurs in Central America and North America. The majority of the genera are Andean-Patagonian. Among the
Perlidae, Kempnyia, Macrogynoplax and Enderleina occur to the east and north in South America (Stark et al.
2009).
Nymphs of these genera are still incompletely studied. Of the fourteen species of Macrogynoplax known, only
four have been associated with the adults. Only one presumed association in Enderleina is known (Stark, 1989). As
with other perlids, adults of Macrogynoplax and Enderleina are often captured using Malaise traps or attracted to
light traps.
There are six species of Macrogynoplax and four of Enderleina described in the Brazilian Amazon and known
primarily from adults. Macrogynoplax is found at high and low altitudes and nymphal habitats are similar to that of
the genus Anacroneuria. Less is known about Enderleina. Recent research has indicated a relatively low
occurrence of these genera in the Central Amazon (Bobot & Hamada, 2002).
Expeditions conducted by the author in the northeastern and southeastern areas of the state of Pará
(microbasins: Ape River, Peixe-Boi River and Serra dos Martírios-Andorinhas) and in the Reserva Floresta
Nacional do Amapá, have indicated that species of these two genera are not common in the eastern Amazon, as in
other states of the Brazilian Amazon (Table 4).
TABLE 4. Collection localities in Brazilian Amazon as mentioned in the text.
State
Municipality
Locality
Latitude
Longitude
Amapá
Macapá
Floresta Nacional do Amapá
0.9717
-51.6139
Amazonas
Manaus
BR174, ZF3
-2.353056
-59.984722
Amazonas
Manaus
Reserva de Desenvolvimento Sustentável do Tupé
-3.0342
-60.2763
Amazonas
Manaus
Reserva Florestal Adolpho Ducke
-2.921689
-59.955203
Pará
Peixe-Boi
Peixe-Boi River Abaeté
-1.139181
-47.323633
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Acknowledgements
The authors thank Dr. C.G. Froehlich (USP) and Dr. Pitágoras da Conceição Bispo (UNESP) for their valuable
suggestions throughout the study. Dr. Augusto Loureiro Henriques (INPA), Dra. Sônia Casari (MZUSP) and Dra.
Inacia Maria Vieira (IEPA) for the loan of specimens, Dr. Cléverson Ranniéri Meira dos Santos (MPEG), Dr.
Emerson Monteiro dos Santos (UFOPA), Dr. Jaime Liege de Gama Neto (UERR), Dr. Bento Melo Mascarenhas
(MPEG) and B.Sc. Cândido Rodrigues Monteiro Júnior (MPEG) for assisting in the collections of material. To Dr.
Gleomar Fabiano Maschio (UFPA), Dra. Sheila Costa Pinheiro (UFPA), B.Sc. Diana Almeida da Silva (UFPA),
B.Sc. Ieda Maria da Silva (UFPA), B.Sc. Maria Lucia da Silva Sales (UFPA), B.Sc. Zilda de Fátima, for their
support in the field work. Technical support was given by Raimundo Nonato Vasques Bittencourt, Domingos
Dalcides Guimarães (MPEG), Luiz Augusto Quaresma (MPEG) and Esmeraldo Cavalcante de Aguiar (MPEG). To
Abel Pojo (SEMA), Adailton Brito (SEMA) and José Lopes da Silva (resident of Serra dos Martírios-Andorinhas),
for their support in work. We thank the Museu Paraense Emílio Goeldi (MPEG); Instituto Nacional de Pesquisas da
Amazônia (INPA); Secretaria de Estado do Meio Ambiente (SEMA); Universidade Federal do Pará (EAD-UFPA);
University of São Paulo (USP); Museu de Zoologia de São Paulo (MZUSP); Instituto de Pesquisas Científica e
Tecnológica do Estado do Amapá (IEPA); Universidade Estadual de Roraima (UERR); Laboratory of Aquatic
Biology of UNESP (Assis, São Paulo); FAPESP (State of São Paulo Research Foundation, grant number 2012/
23058-1), and the National Council of Technological and Scientific Development (CNPq) for institutional and
financial support. We thank two reviewers for improving the manuscript.
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