Zootaxa 3980 (4): 477–500 ISSN 1175-5326 (print edition)

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Copyright © 2015 Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
http://dx.doi.org/10.11646/zootaxa.3980.4.2
http://zoobank.org/urn:lsid:zoobank.org:pub:CDBE7FC5-92D7-4D48-8CB4-FEF1D8587F09

Two new species of Rhagovelia (Hemiptera: Heteroptera: Veliidae)

from Costa Rica, with a key and new records from the country

FELIPE FERRAZ FIGUEIREDO MOREIRA1, 2, BERNALD PACHECO-CHAVES3, 4,

MONIKA SPRINGER3 & ISABELLE DA ROCHA SILVA CORDEIRO1
1
Instituto Oswaldo Cruz, Laboratório Nacional e Internacional de Referência em Taxonomia de Triatomíneos. Avenida Brasil, 4365,
Pavilhão Rocha Lima, Quinto Andar. Manguinhos, Rio de Janeiro, RJ, Brasil. CEP 21045-900. E-mail: felipe.moreira@ioc.fiocruz.br
2
Universidade Federal do Rio de Janeiro, Instituto de Biologia, Departamento de Zoologia, Laboratório de Entomologia. Avenida
Carlos Chagas Filho 373, CCS, Bloco A, Sala 107. Cidade Universitária, Rio de Janeiro, RJ, Brasil. CEP 21941-971, Caixa Postal
68044. E-mail: felipento@hotmail.com
3
Universidad de Costa Rica, Escuela de Biología, Museo de Zoología, CIMAR. San Pedro de Montes de Oca, SJ, Costa Rica. Apar-
tado Postal 11501-2060. E-mails: bernald.pacheco@gmail.com, biomonitoreo@gmail.com
4
Laboratorio de Biomonitoreo Acuático AquaBioLab S. A. San Pedro de Montes de Oca, SJ, Costa Rica. Apartado Postal 2241-2050

Abstract

Rhagovelia danpolhemi sp. nov., from the Caribbean Slope of Costa Rica and R. springerae sp. nov., from the northern
portion of the country, are described, illustrated, compared with similar species, and included in a key to species recorded
from the country. Rhagovelia calopa Drake & Harris, R. spinosa Gould, and R. tantilla Drake & Harris are recorded for
the first time from Costa Rica. New provincial records are provided for R. acuminata Bacon, R. angustipes Uhler, R. ar-
mata (Burmeister), R. bisignata Bacon, R. crassipes Champion, R. elegans Uhler, R. femoralis Champion, R. solida Ba-
con, R. spinigera Champion, R. tenuipes Champion, and R. uncinata Champion. Additional distributional information is
given for these species and for R. formosa Bacon and R. scabra Bacon.

Key words: Aquatic insects, Neotropical Region, taxonomy, water striders

Resumen

Se describe, ilustra, compara con especies similares, y se incluye en una clave para especies registradas en Costa Rica a
Rhagovelia danpolhemi sp. nov., especie de la Vertiente Caribe del país, y R. springerae sp. nov., de la región norte.
Además, se reportan por primera vez para Costa Rica las especies Rhagovelia calopa Drake & Harris, R. spinosa Gould
y R. tantilla Drake & Harris. Se proveen nuevos registros a nivel de provincia para R. acuminata Bacon, R. angustipes
Uhler, R. armata (Burmeister), R. bisignata Bacon, R. crassipes Champion, R. elegans Uhler, R. femoralis Champion, R.
solida Bacon, R. spinigera Champion, R. tenuipes Champion y R. uncinata Champion; y se da información adicional de
la distribución de estas especies y de R. formosa Bacon y R. scabra Bacon.

Palabras clave: Chinches patinadores, insectos acuáticos, región neotropical, taxonomía

Introduction

The members of the broad-shouldered water strider genus Rhagovelia Mayr, 1865 are among the most commonly
encountered aquatic Heteroptera in tropical regions, forming the dominant guild of surface-dwelling insect
predators on rocky upland streams (Polhemus 1997), and Costa Rica is not an exception. Representatives of the
genus can be considered as the most common semiaquatic bugs in streams and rivers from the country, ranging
from lowlands to highlands.
Studies concerning this genus in Costa Rica are scarce, although a new species was recently described

Accepted by M. Malipatil: 9 Jun. 2015; published: 1 Jul. 2015 477

(Buzzetti and Zettel 2007) from a locality with a high diversity of Rhagovelia, where ten species were collected in
an area of only 6 km2 (Hecher and Zettel 1996). This paper is the result of several years of field collections in Costa
Rica, and the study of specimens of Rhagovelia in the Collection of Aquatic Entomology of the Museo de
Zoología, Universidad de Costa Rica.

Material and methods

Type material has been deposited on the Museo de Zoología, Escuela de Biología, Universidad de Costa Rica
(MZUCR) and the Coleção Entomológica Professor José Alfredo Pinheiro Dutra, Laboratório de Entomologia,
Departamento de Zoologia, Instituto de Biologia, Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil
(DZRJ). All other material is deposited in MZUCR. Measurements are given in millimeters and abbreviated as
follows: body length (BL), head length (HL), head width through eyes (HW), length of antennomeres I–IV
[without intersegmental pieces] (ANT I, ANT II, ANT III, ANT IV), maximum eye width (EYE), pronotum length
on midline (PL), greatest pronotum width (PW), femoral length (FEM), tibial length (TIB), and length of
tarsomeres I–III (TAR I, TAR II, TAR III). Descriptions and photographs were made using specimens preserved in
alcohol.

Results and discussion

Descriptions

Rhagovelia danpolhemi Moreira, Pacheco-Chaves & Cordeiro sp. nov.

(Figs 1–8, 28)

Apterous male (Fig. 1, Table 1). General color very dark brown to black, covered by silvery pubescence on head
and body. Head with impressed longitudinal midline and pair of indentations on base shining black. Eyes dark red.
Antenniferous tubercles dark brown. Buccula and base of rostrum dark brown; rostrum shining, becoming black
apically. Antenna with base of article I white to pale yellow and remaining brown to dark brown, lighter on
intersegmental pieces. Pronotum with transverse, centrally-interrupted, orange stripe between eyes. Mesonotum
anteriorly with two oblique shining black areas. Proacetabulum with most of venter and margin yellowish brown;
mesoacetabulum with margin yellowish brown; metacetabulum with posterior half yellowish brown. Abdominal
tergites VI–VII centrally shining black, sometimes also with small central mark on tergite V; venter entirely dark
bluish brown, or bluish black with central subquadrate area of last sternite dark brown to black. Genital segments
shining dark brown to black. Fore and hind coxae yellow to yellowish brown; mid coxa brown to dark brown; fore
trochanter light brown to black; mid trochanter dark brown with base sometimes lighter; hind trochanter yellow to
yellowish brown; fore and hind femora entirely dark brown or with a small light brown area on base contrasting
with rest of segment. Remainder of legs dark brown to black.
Head and body velvety. Head short, with a few long setae anteriorly to eyes and adjacent to inner eye margin;
without denticles. Antenna covered by short brown setae, with additional stout erect setae on articles I–II.
Antennomeres I–III cylindrical; IV fusiform; I thicker than others, arcuate, curving laterad; III slightly broader at
apex than II and IV. Rostrum thick, reaching base of mesosternum.
Thorax dorsally covered by short, shining setae and longer light setae on posterior margin of meso- and
metanota; laterally with long, dark, curved setae. Pronotum short, centrally flat, with posterior margin nearly
straight, not covering mesonotum. Mesonotum slightly swollen centrally. Venter of thorax without denticles or
other distinct features.
Abdomen dorsally covered by short, golden setae. Abdominal tergites I–V short and wide; VI longer; VII
subquadrate. Posterior margins of abdominal tergites I–III slightly convex; of tergites IV–VII straight or nearly so.
Connexiva horizontal to slightly elevated, subparallel from base to tergite IV, then tapering to apex. Sternites with
posterior margins concave. Last sternite distinctly longer than others. Proctiger as in Fig. 2. Paramere as in Fig. 3.

478 · Zootaxa 3980 (4) © 2015 Magnolia Press MOREIRA ET AL.

FIGURES 1–8. Rhagovelia danpolhemi sp. nov. 1. Apterous male, dorsal view. 2. Male proctiger. 3. Male paramere, lateral
view. 4. Detail of male left fore leg, ventral view. 5. Apex of male abdomen and left hind leg, ventral view. 6. Apterous female,
dorsal view. 7. Detail of female left fore leg, ventral view. 8. Apex of female abdomen and hind legs, ventral view. Scale bar =
1.00 mm.

TWO NEW SPECIES OF RHAGOVELIA Zootaxa 3980 (4) © 2015 Magnolia Press · 479
 Legs covered by short, brown setae and rows of longer, thicker setae on femora and tibiae. All legs without
spines or denticles. Fore femur slightly arcuate. Fore tibia more strongly arcuate (Fig. 4), slightly concave near
apex of inner surface. Hind femur elongate, distinctly thinner than base of mid femur. Hind tibia without apical
spur (Fig. 5).

TABLE 1. Descriptive measurements of apterous males of Rhagovelia danpolhemi sp. nov.

Structure Specimen 1 Specimen 2 Specimen 3 Specimen 4 Specimen 5
BL 2.36 2.44 2.48 2.52 2.36
HL 0.29 0.30 0.34 0.39 0.31
HW 0.81 0.83 0.84 0.84 0.78
ANT I 0.78 0.82 0.97 0.87 0.86
ANT II 0.46 0.42 0.50 0.53 0.49
ANT III 0.48 0.50 0.59 0.56 0.53
ANT IV 0.48 0.50 0.51 0.53 0.49
EYE 0.27 0.29 0.29 0.23 0.26
PL 0.16 0.16 0.19 0.17 0.20
PW 0.75 0.84 0.81 0.86 0.91
FEM 1.00 1.14 1.00 1.06 1.06
TIB 1.04 1.20 1.12 1.08 1.12
TAR I 0.03 0.03 0.03 0.03 0.03
TAR II 0.03 0.03 0.03 0.03 0.03
TAR III 0.23 0.26 0.23 0.27 0.24
FEM 1.73 1.87 1.83 1.90 1.80
TIB 1.17 1.27 1.27 1.13 1.20
TAR I 0.11 0.07 0.09 0.07 0.07
TAR II 0.64 0.77 0.71 0.71 0.73
TAR III 0.71 0.79 0.79 0.74 0.76
FEM 1.06 1.33 1.30 1.33 1.25
TIB 1.46 1.80 1.75 1.63 1.63
TAR I 0.04 0.06 0.06 0.06 0.06
TAR II 0.07 0.09 0.10 0.10 0.09
TAR III 0.21 0.27 0.27 0.29 0.27

Macropterous male (Table 2). Identical to apterous male except for structure of thorax and wing
development. Pronotum subpentagonal, with posterior lobe developed and completely covering remainder of
thorax; posterior corner rounded, without projections. Wings broken off approximately at middle. Forewings with
two elongated proximal cells and two small rounded distal cells. Outer vein of forewing densely covered by brown
setae, especially distally.
Apterous female (Fig. 6, Table 3). Similar in color and structure to apterous male, but larger and more robust.
Light areas of acetabula sometimes more extensive and paler than in males. Fore and hind coxae pale yellow to
yellowish brown; mid coxa brown to black; fore and hind trochanters pale yellow to yellowish brown; mid
trochanter dark brown with base sometimes lighter; fore femur yellow to yellowish brown from base almost to
middle, then dark brown; hind femur entirely dark brown or with a small light brown area on base contrasting with
rest of segment. Remainder of legs dark brown to black. Shining black areas on abdominal tergites V–VIII.
Connexiva horizontal to slightly elevated. Legs without spines, denticles or spurs. Fore femur and tibia less curved
than in male (Fig. 7). Hind femur elongated, thinner than middle femur at base (Fig. 8).
Macropterous female (Table 4). Identical to apterous female, except structure of thorax and wing
development, which are like those of macropterous males.

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TABLE 2. Descriptive measurements of macropterous males of Rhagovelia danpolhemi sp. nov.
Structure Specimen 1 Specimen 2 Specimen 3 Specimen 4 Specimen 5
BL 2.52 2.44 2.48 2.60 2.40
HL 0.30 0.27 0.33 0.31 0.24
HW 0.80 0.81 0.81 0.89 0.84
ANT I 0.91 0.89 0.89 0.91 0.86
ANT II 0.50 0.46 0.49 0.49 0.49
ANT III 0.54 0.54 0.53 0.57 0.51
ANT IV 0.46 0.49 0.50 0.54 0.53
EYE 0.27 0.29 0.29 0.29 0.27
PL 1.29 1.24 1.30 1.31 1.26
PW 1.37 1.40 1.39 1.44 1.30
FEM 1.06 1.02 1.04 1.02 0.98
TIB 1.12 1.10 1.12 1.10 1.04
TAR I 0.03 0.03 0.03 0.03 0.03
TAR II 0.03 0.03 0.03 0.03 0.03
TAR III 0.23 0.23 0.23 0.27 0.23
FEM 1.90 1.83 1.77 1.87 1.70
TIB 1.13 1.13 1.20 1.17 1.10
TAR I 0.07 0.07 0.07 0.06 0.06
TAR II 0.64 0.63 0.64 0.61 0.61
TAR III 0.67 0.74 0.70 0.70 0.73
FEM 1.25 1.33 1.35 1.33 1.33
TIB 1.68 1.58 1.60 1.60 1.55
TAR I 0.06 0.06 0.06 0.06 0.06
TAR II 0.08 0.10 0.09 0.10 0.10
TAR III 0.26 0.30 0.29 0.29 0.30

Type material. COSTA RICA: Heredia—Sarapiquí, Sarapiquí River, 200–275 m a.s.l., III.1998, (P. Paaby): 1
apterous male [HOLOTYPE], 3 apterous males, 5 apterous females [PARATYPES] (DZRJ). Sarapiquí,
Tirimbina Road, Sarapiquí River, 03.X.2005, (M. Werner): 3 apterous males [PARATYPES] (MZUCR).
Sarapiquí, Horquetas, Puerto Viejo River, Hydropower Project Cubujuquí, 18.III.2012, (B. Pacheco): 1 apterous
male, 3 apterous females [PARATYPES] (DZRJ). Sarapiquí, InBio Station, Costa Rica River, 05.XI.2002, (A. Y.
Jiménez): 1 male with broken wings [PARATYPE] (DZRJ). Sarapiquí, 17.III.2006, (B. Pacheco): 1 apterous
female [PARATYPE] (MZUCR). Cariblanco, ICE Hydropower Project, 05.VII.2012, (B. Pacheco & F. Reyes): 1
apterous male, 1 male with broken wings, 3 apterous females, 1 female with broken wings [PARATYPES]
(DZRJ). Cariblanco, Cariblanco River, 05.VII.2012, (B. Pacheco & F. Reyes): 1 male with broken wings
[PARATYPE] (MZUCR). Alajuela—San Carlos, Balsa River, Hydropower Project before Tapesco, 04.VIII.2011,
(R. Lara): 1 male with broken wings, 1 apterous female [PARATYPES] (MZUCR). San Carlos, Venecia, Hidalgo
Stream, 10o22.515’ N / 84o16.955’, 338 m a.s.l., (B. Pacheco): 1 apterous female [PARATYPE] (MZUCR). San
Carlos, Florencia, Río La Vieja, power house of Hydropower Project Chocosuela II, 04.XII.2011, (M. Springer): 2
apterous males, 1 apterous female [PARATYPES] (MZUCR). San Carlos, Aguas Zarcas, Negritos River,
10o22.381’ N / 84o19.270’ W, 446 m a.s.l., 23.VIII.2008, (B. Pacheco): 2 apterous males [PARATYPES]
(MZUCR). San Ramón, Balsa River, 400 m a.s.l., 27.IV.2002, (M. Mug): 1 apterous male [PARATYPE] (DZRJ).
San Ramón, San Ramón Reserve, San Lorencito River, 860 m a.s.l., 16.II.1994, (M. Springer): 1 male with broken
wings [PARATYPE] (DZRJ) San Ramón, San Lorenzo River, (B. Pacheco & F. Reyes): 1 apterous male
[PARATYPES] (MZUCR). San Ramón, hydropower station, San Lorenzo, above dam, 300 m a.s.l., 20.V.2011,

TWO NEW SPECIES OF RHAGOVELIA Zootaxa 3980 (4) © 2015 Magnolia Press · 481
(D. Vásquez): 4 apterous males, 6 apterous females [PARATYPES] (MZUCR); above dam, 330 m a.s.l.,
20.V.2011, (B. Pacheco): 12 apterous males, 1 macropterous male, 17 apterous females, 1 macropterous female
[PARATYPES] (MZUCR). Cartago—Paraíso, Orosi River, 20.IX.1998, (X. Miranda): 1 apterous female
[PARATYPE] (MZUCR); Orosi River, 1400 m a.s.l., 20.IX.1998, (P. Ortiz): 1 apterous female [PARATYPE]
(DZRJ). Paraíso, Orosi, Quirí River, 1157 m a.s.l., 28.VIII.2005, (G. Vargas): 1 apterous male [PARATYPE]
(MZUCR). Tapantí, Orosi River, 23.X.2005 (M. Werner): 5 apterous males, 9 apterous females [PARATYPES]
(MZUCR). Turrialba, Atirro River, Reventazón Project, 586 m a.s.l., 25.VI.2011, (B. Pacheco & F. Reyes): 1
apterous male, 3 apterous females [PARATYPES] (DZRJ). Turrialba, Jícotea River, 780 m a.s.l., 15.VII.1991, (M.
Springer): 1 apterous female [PARATYPE] (MZUCR). Limón—Pococí, Guápiles, Blanco River, strong current,
235 m a.s.l., 14.XI.2005, (C. Lizana): 1 apterous male, 2 apterous females [PARATYPES] (MZUCR); Blanco
River, 100 m a.s.l., 17.XI.2005, (E. Boza): 1 apterous male [PARATYPE] (DZRJ). Pococí, Guápiles, Costa Rica
River, under the bridge, 232 m a.s.l, 14.XI.2005, (C. Lizana): 1 apterous male, 1 apterous female [PARATYPE]
(MZUCR). Guápiles, Chirripó River, site 1, above CDP, 07.IV.2011, (B. Pacheco): 5 apterous males, 1
macropterous male, 6 apterous females, 1 macropterous female [PARATYPES] (MZUCR); Chirripó River, site 2,
inside CDP: 7 apterous males, 3 macropterous males, 13 apterous females, 2 macropterous females
[PARATYPES] (MZUCR).

TABLE 3. Descriptive measurements of apterous females of Rhagovelia danpolhemi sp. nov.

Structure Specimen 1 Specimen 2 Specimen 3 Specimen 4 Specimen 5
BL 2.72 2.92 2.92 2.84 2.88
HL 0.29 0.33 0.29 0.37 0.35
HW 0.89 0.90 0.89 0.87 0.87
ANT I 0.86 0.93 0.83 0.91 0.84
ANT II 0.49 0.49 0.49 0.47 0.47
ANT III 0.53 0.59 0.55 0.50 0.50
ANT IV 0.50 0.51 0.52 0.47 0.49
EYE 0.24 0.27 0.27 0.30 0.27
PL 0.16 0.21 0.21 0.21 0.21
PW 0.86 0.90 0.87 0.99 0.91
FEM 0.96 1.04 1.08 0.92 0.94
TIB 0.98 1.06 1.10 0.94 1.00
TAR I 0.03 0.04 0.04 0.04 0.04
TAR II 0.03 0.04 0.04 0.04 0.04
TAR III 0.29 0.27 0.30 0.27 0.27
FEM 1.70 1.97 1.90 1.77 1.73
TIB 1.17 1.30 1.27 1.13 1.20
TAR I 0.09 0.07 0.07 0.07 0.09
TAR II 0.70 0.74 0.84 0.64 0.66
TAR III 0.79 0.83 0.86 0.79 0.81
FEM 1.24 1.35 1.35 1.18 1.28
TIB 1.42 1.60 1.63 1.53 1.48
TAR I 0.06 0.06 0.05 0.07 0.07
TAR II 0.11 0.10 0.11 0.11 0.11
TAR III 0.30 0.33 0.33 0.31 0.29

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TABLE 4. Descriptive measurements of the macropterous female of Rhagovelia danpolhemi sp. nov.
Structure Specimen 1
BL 3.04
HL 0.34
HW 0.87
ANT I 0.86
ANT II 0.44
ANT III 0.47
ANT IV 0.46
EYE 0.27
PL 1.41
PW 1.53
FEM 1.02
TIB 1.04
TAR I 0.04
TAR II 0.04
TAR III 0.29
FEM 1.70
TIB 1.17
TAR I 0.07
TAR II 0.63
TAR III 0.69
FEM 1.28
TIB 1.50
TAR I 0.06
TAR II 0.11
TAR III 0.31

Distribution. So far the species has been collected only on the Caribbean Slope of Costa Rica (Fig. 28).
Despite of all collecting efforts made on the Pacific Slope, this species has not been found in the area.
Etymology. Named in honor of Dr. Dan A. Polhemus for his great contributions to the taxonomy of the genus
Rhagovelia.
Comments. Rhagovelia danpolhemi sp. nov. belongs to the angustipes complex sensu Polhemus (1997),
based on apterous forms having the pronotal midline shorter than the dorsal length of an eye. Because
macropterous specimens are available, which is uncommon in the complex, placement of the species in the
bisignata group is possible due to the presence of four closed cells on the forewings, including two elongated
proximal cells and two short distal cells. The wings of the macropterous specimens we observed are broken off
near the middle; however, the cells are visible in all specimens.
The absence of spines on the relatively thin hind femur is a feature common in females of the angustipes
complex; however, only five species have unarmed hind femora in both females and males: R. festae Kirkaldy,
1899, R. imitatrix Bacon, 1948, R. longipes Gould, 1931, R. magdalena Padilla-Gil, 2011, and R. danpolhemi sp.
nov. In addition to the main differences among these species (Table 5), the shape of the male parameres also differs
between them (see Bacon 1956, Padilla-Gil 2011).

TWO NEW SPECIES OF RHAGOVELIA Zootaxa 3980 (4) © 2015 Magnolia Press · 483
 TABLE 5. Structural features of the five species of the angustipes complex with unarmed hind femur in both males and females.

Structure R. festae R. imitatrix R. longipes R. magdalena R. danpolhemi

BL apterous male 2.60 3.40 3.00 3.84 2.36–2.52

BL macropterous male ? 3.80 3.05 3.88 2.40–2.60

BL apterous female 2.60 4.06 3.50 4.66 2.72–2.92

BL macropterous female ? 4.32 3.45 4.80 3.04

484 · Zootaxa 3980 (4) © 2015 Magnolia Press

ANT II/ANT III 1.33 1.00 0.70–0.71 0.81-0.85 0.83–0.96

Hind femur/tibia male 0.90–1.00 0.69 0.64 0.58 0.72–0.86

Hind femur/tibia female 0.90–1.00 0.77 0.73 ? 0.77–0.87

Fore coxa brownish yellow dark brown to black, rarely brown to yellow black pale yellow to yellowish brown

Fore trochanter bluish-black black dark brown to black, rarely brown to yellow black pale yellow to dark brown

Mid coxa brownish yellow dark brown to black black brown to black

Mid trochanter bluish-black black dark brown to black black dark brown, sometimes lighter on base

Hind coxa brownish yellow dark brown to black, rarely brown to yellow yellow pale yellow to yellowish brown

Hind trochanter bluish-black black dark brown to black, rarely brown to yellow yellow pale yellow to yellowish brown

Shining tergites male ? at most VII at most VII VII VI–VII, rarely V–VII

Shining tergites female ? at most VIII at most VIII VIII V–VIII

MOREIRA ET AL.
Rhagovelia springerae Moreira, Pacheco-Chaves & Cordeiro sp. nov.
(Figs. 9–14, 29)

Apterous male (Fig. 9, Table 6). Head black. Antenniferous tubercles, clypeus and eyes brown. Antennomere I
with basal ca. half pale yellow; remainder of antenna brown, except yellow intersegmental pieces. Buccula yellow.
Rostrum with article I yellowish brown, II orange-brown, III dark brown, IV black. Pronotum black with two
orange patches anteriorly and circular punctations throughout length. Sides and venter of thorax grayish black.
Abdominal tergites black. Connexiva black with orange-brown on lateral margin. Abdominal sternites grayish
black. Genitalia brown, covered by setae dorsally; ventrally orange-brown. All coxae and trochanters yellow. Fore
femur yellow basally, dark brown apically; mid femur dark brown dorsally, brown ventrally; hind femur dark
brown, lighter basally. Fore tibia dark brown dorsally, brown ventrally; mid and hind tibiae and all tarsi dark
brown.

TABLE 6. Descriptive measurements of apterous males of Rhagovelia springerae sp. nov.

Structure Specimen 1 Specimen 2
BL 3.03 (genitalia removed) 3.50 (genitalia not removed)
HL 0.39 0.36
HW 0.84 0.87
ANT I 0.75 0.81
ANT II 0.45 0.45
ANT III - 0.42
ANT IV - 0.45
EYE 0.27 0.30
PL 0.96 0.96
PW 1.15 1.15
FEM 0.90 0.90
TIB 0.93 0.87
TAR I 0.05 0.04
TAR II 0.04 0.05
TAR III 0.21 0.28
FEM 1.57 1.51
TIB 1.12 1.09
TAR I 0.08 0.07
TAR II 0.52 0.53
TAR III 0.74 0.73
FEM 1.39 1.40
TIB 1.21 1.18
TAR I 0.06 0.05
TAR II 0.13 0.11
TAR III 0.34 0.34

Head short, wide, velvety, covered by short brown setae and longer, thicker, black setae. Antenna covered by
short setae, articles I–II also with long setae. Antennomere I curved outside, narrow on base, stout on apex; II–III
cylindrical, IV fusiform. Jugum, proepisternum and proacetabulae with black denticles (Fig. 10). Rostrum reaching
middle of mesosternum.
Pronotum completely covering mesonotum, with longitudinal median carina weakly developed (better
observed on wet specimen). Mesosternum with two oblique carinae covered by long setae. Meso- and metasterna
with sparse black denticles, obscured by the dark body color.

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FIGURES 9–14. Rhagovelia springerae sp. nov. 9. Apterous male, dorsal view. 10. Male head and part of thorax, ventral view,
white arrow shows area with black denticles. 11. Male abdomen, ventral view. 12. Male proctiger. 13. Male paramere, lateral
view. 14. Male left hind leg, ventral view. Scale bar = 1.00 mm.

486 · Zootaxa 3980 (4) © 2015 Magnolia Press MOREIRA ET AL.

 Abdominal tergites and sternites covered by brown setae; sternites also with black denticles in center, obscured
by the dark body color. Longitudinal median carina on visible sternites I–III, weak on III (Fig. 11). Deep depression
centrally on visible sternites V–VII, sharply on VI–VII (Fig. 11). Robust black denticles absent from posterolateral
margins of last abdominal segment surrounding genital cavity. Proctiger as in Fig. 12. Parameres symmetrical,
shape as in Fig. 13.
Legs covered by short, brown setae, with erect, black setae on femora and tibiae. Coxae granulose. Venter of
fore trochanter distally with tuft of setae (Fig. 10). Fore tibia widening apically. Mid femur narrowing apically.
Hind trochanter unarmed (Fig. 11). Hind femur stout, without spines on proximal 1/3; distal 2/3 with dorsal row of
9–11 spines decreasing in size (Fig. 14) and ventral row of 4–5 short spines. Hind tibia with two irregular rows of
short spines, two slightly displaced spines close to apex, and a straight apical spur (Fig. 14).
Type material. COSTA RICA: Alajuela—Upala, Canalete, Achiote River, desfogue, VIII.2010, (F. Reyes &
P. Gutiérrez): 1 apterous male [HOLOTYPE] (MZUCR). Guanacaste—Cañas, site 2, below confluence, dam,
EsIA south channel, 47 m a.s.l., VII.2010, (B. Pacheco & D. Vásquez): 1 apterous male [PARATYPE] (DZRJ).
Distribution. Despite the extensive collecting effort in Costa Rica, only two representatives of this species
have been found, both collected in the northern portion of the country (Fig. 29).
Etymology. Named in honor of MSc. Monika Springer for her great contributions to the aquatic entomology in
Costa Rica.
Comments. Rhagovelia springerae sp. nov. belongs to the robusta group sensu Polhemus (1997), based on the
pronotum of apterous forms completely covering the mesonotum, the posterolateral margins of the last abdominal
segment surrounding the genital cavity without robust black spines, the hind tibia with a straight apical spur, and by
the shape of the male parameres.
In the key to the species of the group provided by Polhemus (1997), R. springerae sp. nov. runs to R. zeteki
Drake, 1953, known from Colombia and Panama. However, the new species can be differentiated from it by the
brownish coloration of the lateral margins of the connexiva; central depression on male visible abdominal sternites
V–VII; the tuft of long, stout setae on the distal portion of the fore trochanter; the absence of small spines on the
basal 1/3 of hind femur; and by the shape of the male parameres and proctiger. Rhagovelia springerae sp. nov. also
can be distinguished from other species of the group known from Costa Rica by the characters mentioned in the
key below.

Key to the Rhagovelia from Costa Rica [modified from Bacon (1956) and Polhemus (1997)]

1 Pronotum of apterous forms at midline shorter than dorsal length of an eye (Fig. 15) . . . . . . . . . . . . (angustipes complex)... 2
- Pronotum of apterous forms at midline longer than dorsal length of an eye (Fig. 16) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
2 Hind femur unarmed in both males and females. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhagovelia danpolhemi sp. nov.
- Hind femur with spines at least in males. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Base of last abdominal sternite of male bearing a stout, slightly curved spine. . . . . . . . . . . . . . . . . . . . . . . . Rhagovelia spinosa
- Base of last abdominal sternite of male without such a spine. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 Hind trochanter of male and female with short spines; hind femur of male strongly incrassate . . . . . . . . . . Rhagovelia calopa
- Hind trochanter of male and female without spines; hind femur of male not strongly incrassate . . . . . . . . . . . . . . . . . . . . . . . 5
5 Antennomere II longer than or subequal to III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
- Antennomere II shorter than III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
6 Hind tibia of male and female with short spines along entire length (those of females may be weak, but present), besides apical
spur . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhagovelia tenuipes (part)
- Hind tibia of male and female without short spines along its length, at most with apical spur . . . . . . . . . Rhagovelia bisignata
7 Hind tibia of male and female with short spines along entire length (those of females may be weak, but present), besides apical
spur . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhagovelia tenuipes (part)
- Hind tibia of male and female without short spines along its length, at most with apical spur . . . . . . . . . . . . . . . . . . . . . . . . . 8
8 Male hind tibia without apical spur; apterous female with shining black spots on last two abdominal tergites . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhagovelia tantilla
- Male hind tibia with apical spur; apterous female with shining black spots on last four or five abdominal tergites . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Rhagovelia angustipes
9 Male with posterolateral margins of abdominal segment VII surrounding genital cavity bearing large black denticles (Fig. 17);
macropterous female with an elevated posterior projection at apex of pronotum (Fig. 18A, B) . . . . . . . . . . . . . . . . . . . . . . . 10
- Male with posterolateral margins of abdominal segment VII surrounding genital cavity without large black denticles;
macropterous female without elevated posterior projection at apex of pronotum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
10 Female with central section of middle femur with a transverse constriction (Fig. 19). . . . . . . . . . . . . . . . . (spinigera group) 11

TWO NEW SPECIES OF RHAGOVELIA Zootaxa 3980 (4) © 2015 Magnolia Press · 487
- Female with central section of middle femur without a transverse constriction. . . . . . . . . . . . . . . . . . . . . . . . (armata group) 12
11 Male middle femur armed with numerous small black spines on proximal 1/3 of posterior margin; female with long, slender,
erect black setae often present on margins of connexiva VI–VII (most easily seen in lateral view) . . . . . Rhagovelia spinigera
- Male middle femur lacking small black spines on proximal 1/3 of posterior margin; female lacking erect, black setae on mar-
gins of connexiva VI–VII . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhagovelia formosa
12 Male and female with terminal genital segment acuminate, prolonged into a spine-like process (similar to Fig. 20). . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhagovelia acuminata
- Male and female terminal genital segment not acuminate, sometimes ending in an acute angle, but never prolonged into a
spine-like process (similar to Fig. 21) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13 Female with connexival margins relatively straight, tapering evenly toward apex of abdomen, never suddenly convergent adja-
cent to or after tergite III (similar to Fig. 22) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhagovelia crassipes
- Female with connexival margins sharply convergent adjacent to or after abdominal tergite III (similar to Fig. 23). . . . . . . . 14
14 Lateral portions of mesosternum lacking small black denticles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhagovelia armata
- Lateral portions of mesosternum bearing numerous small black denticles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15 Disk of pronotum thickly covered by moderately long, recumbent pale pubescence; venter of abdomen with numerous small
black denticles along its entire length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhagovelia scabra
- Disk of pronotum moderately covered by short, recumbent golden setae; venter of abdomen with black denticles on lateral por-
tions of basal segments only . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhagovelia solida
16 Male and female hind tibia with crescent-shaped apical spur (Fig. 24) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (elegans group) 17
- Male and female hind tibia with apical spur, when present, straight or slightly curved (Fig. 14) . . . . . . . . . . . . . . . . . . . . . . 18
17 Male and female with terminal genital segment acuminate posteriorly, prolonged into an extremely slender spine-like process
(Fig. 20). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhagovelia uncinata
- Male and female terminal genital segment not acuminate posteriorly, often ending in an acute angle, but not prolonged into a
spine-like process (Fig. 21) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhagovelia elegans
18 Abdomen of apterous female with dorsal longitudinal median carina (indistinct in some cases) (Fig. 25); abdominal connexiva
of apterous female thickened in some species (Fig. 25); male hind tibia only with subequal spines, besides apical spur. . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (hirtipes group) 19
- Abdomen of apterous female without longitudinal carina; abdominal connexiva of apterous female never thickened; male hind
tibia of some species displaying one or more spines larger than the others, besides apical spur (as in Fig. 26) . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (robusta group) 20
19 Apterous female connexiva bowed inward and glabrous adjacent to abdominal tergites III–V; male abdominal sternite VIII
lacking a median carina. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhagovelia reclusa
- Apterous female connexiva not bowed inward and glabrous adjacent to tergites III–V; male abdominal sternite VIII usually
with a low median carina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhagovelia femoralis
20 Length of apterous forms 3.90–4.10 mm; male hind trochanter bearing 2–4 small subequal spines; male hind tibia with small
peg-like spines throughout most of its length and slightly enlarged spines approximately on 1/3 distance to apex . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhagovelia johnpolhemi
- Length of apterous forms at most 3.50 mm; male hind trochanter bearing 0–1 small spine; male hind tibia with small subequal
peg-like spines. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
21 Male fore trochanter without tuft of long stout setae; male hind femur weakly widened; male parameres with distal portion
long, slender and curved (Fig. 27). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhagovelia sehnali
- Male fore trochanter with a tuft of long stout black setae (Fig. 10); male hind femur moderately widened; male parameres with
distal portion not elongated and curved (Fig. 13) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhagovelia springerae sp. nov.

New records

Rhagovelia acuminata Bacon, 1956

(Fig. 30)

Known distribution in Costa Rica. Puntarenas (Polhemus 1997). Reported here are the first records from
Alajuela and Cartago provinces.
Material examined. COSTA RICA: Alajuela—San Ramón, Alberto Manuel Brenes Biological Reserve, 800
m a.s.l., 10.X.2010, (B. Pacheco): 5 apterous males, 4 apterous females, 2 nymphs; Alberto Manuel Brenes
Biological Reserve, San Lorencito River, 10o13’6.8” N / 84o35’48.9” W, 876 m a.s.l., 10.X.1998: 4 apterous males,
3 apterous females; 17.X.2009, (B. Pacheco): 1 apterous male, 1 apterous female. Cartago—Paraíso, Birrisito,
Parruas River Stream, secondary Forest, 1350 m a.s.l., 15.X.1996, (C. Guillén): 3 apterous males, 4 apterous
females. Puntarenas—Chomes, 04.XII.1998: 2 apterous females. Santa Elena, Berros Stream, 1200 m a.s.l.,
IV.2001, (A. Burgin): 6 apterous males, 6 apterous females.

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FIGURES 15–27. 15. Rhagovelia tenuipes, apterous form, head and thorax, dorsal view. 16. R. elegans, apterous form, head
and thorax, dorsal view. 17. R. scitula, apex of male abdomen with genitalia removed, posterior view. 18–19. R. formosa. 18.
Macropterous female pronotum: A. Dorsal view, B. Lateral view. 19. Female middle femur, dorsal view. 20. R. uncinata, male
proctiger. 21. R. elegans, male proctiger. 22. R. perfecta, apterous female, dorsal view with legs omitted. 23. R. armata,
apterous female, dorsal view with legs omitted. 24. R. elegans, detail of apex of hind tibia with crescent-shaped spur. 25. R.
tijuca, apterous female, dorsal view with legs omitted. 26. R. boliviana, male hind leg, ventral view. 27. R. sehnali, male
paramere [DEN = denticles, Ms = mesonotum, Mt = metanotum, P = pronotum; 18–23 and 25–26 modified from Polhemus
(1997); 24 modified from Moreira and Ribeiro (2009); 27 modified from Buzzetti & Zettel (2007)].

TWO NEW SPECIES OF RHAGOVELIA Zootaxa 3980 (4) © 2015 Magnolia Press · 489
Rhagovelia angustipes Uhler, 1894
(Fig. 28)

Known distribution in Costa Rica. San José (Hungerford 1939). Reported here are the first records from
Alajuela, Cartago, Guanacaste, Heredia, and Puntarenas provinces.
Material examined. COSTA RICA: Alajuela—San Ramón, Las Musas, 2500 m a.s.l., 22.IX.2003, (J. A.
Vindas): 1 apterous male, 1 nymph. Cartago—Tapantí, Orosi River, 23.X.2005, (M. Werner): 1 apterous male, 2
apterous females. Guanacaste—Palo Verde, Lomas Barbudal Corridor, Barbudal Stream, 15.I.2007, (B. Pacheco):
1 apterous male. Bagaces, Lomas Barbudal, river, 300 m a.s.l., 04.X.1998, (P. Ortiz): 1 apterous male. Heredia—
Sarapiquí, Puerto Viejo, La Selva Biological Station, El Surá River, under 50 m a.s.l., 27.VIII.2005, (M. Losilla): 2
apterous females. Sarapiquí, Selva Verde Lodge, Sarapiquí River, 100 m a.s.l., 30.IV.2000, (M. Springer): 1
apterous male, 1 macropterous male, 3 apterous females. Puntarenas—Cuenca del Térraba, Diquís Hydropower
Project, Convento River, 25.II.2005, (R. Lara): 1 apterous male; Diquís Hydropower Project, Brujo River,
01.III.2006, (R. Lara): 1 apterous male; Diquís Hydropower Project, Unión River, 14.III.2005 (R. Lara): 3 apterous
males; Diquís Hydropower Project, ICE Volcán River, 13.II.2006, (R. Lara): 1 apterous male, 2 apterous females.
Quepos, Cañas River, 05.X.2009, (B. Pacheco): 1 apterous female. Osa, Osa Peninsula, Rincón River, under 50 m
a.s.l., 20.VII.1996, (G. Umaña): 1 apterous female. Osa, Tigre River, by the bridge, 27.IX.1996, (G. Umaña): 1
apterous male. Osa, Corcovado National Park, Claro River, near Sirena, 0 m a.s.l., 24.IV.1999, (L. Stauffer): 1
macropterous male, 1 apterous female. Puntarenas, Lepanto, Blanco River, Paso Tres Ríos, 350 m a.s.l., 23.I.1998:
1 apterous male, 2 macropterous males, 3 males with broken wings, 1 female with broken wings. Golfito, Cañaza
Stream, bridge leaving Golfito, 0 m a.s.l., I.1997, (A. Villalobos): 1 apterous male. Golfito, tributary to Cañaza
Stream, I.1997, (A. Villalobos): 8 apterous males, 3 males with broken wings, 9 apterous females. Golfito, La
Gamba Stream, 2 km from Barrio Ureña, primary forest, 120 m a.s.l., I.1997: 2 apterous males, 1 apterous female.
Buenos Aires, Longo Mai, Sonador River, 600 m a.s.l., 23.V.1998, (A. Nunn): 1 apterous male. Coto Brus, Las
Cruces, Jaba River (Matadero), 10.X.1999, (M. Springer): 1 apterous male.

Rhagovelia armata (Burmeister, 1835)

(Fig. 30)

Known distribution in Costa Rica. Cartago, Limón, San José, and Río Virilla [unknown province] (Bacon 1956,
Polhemus 1997). Reported here are the first records from Alajuela and Puntarenas provinces.
Material examined. COSTA RICA: Alajuela—San Carlos, San Pedro (La Tigra), unnamed stream, tributary
of Peñas Blancas River, 150 m a.s.l., 20.X.2001, (M. Springer & C. Guillén): 1 apterous male, 1 apterous female, 1
nymph. San Carlos, Ciudad Quesada, San Vicente, Peje River, 1500 m a.s.l., 14.VIII.2005, (P. Benavides): 4
macropterous males, 2 apterous females. San Ramón, La Paz River: 2 apterous females. San Ramón, San Juan
River, 950 m a.s.l., 7.XII.2003, (N. Zuñiga): 1 apterous female. Cartago—Orosi, 22.XI.2008, (B. Pacheco): 2
apterous females. Orosi, Tapantí National Park, unnamed small stream, 1400 m a.s.l., 13.IX.1996: 1 apterous male,
1 apterous female, 1 macropterous female, 16 nymphs. Puntarenas—Cuenca del Térraba, Diquís Hydropower
Project, Unión River, 14.III.2006, (R. Lara): 1 apterous female.

Rhagovelia bisignata Bacon, 1948

(Fig. 31)

Known distribution in Costa Rica. San José (Bacon 1956). Reported here is the first record from Alajuela
Province.
Material examined. COSTA RICA: Alajuela—Upala, Canalete, Achiote River, desfogue, VIII.2010, (F.
Reyes & P. Gutiérrez): 1 apterous female.

490 · Zootaxa 3980 (4) © 2015 Magnolia Press MOREIRA ET AL.

Rhagovelia calopa Drake & Harris, 1927
(Fig. 28)

Previously recorded from Mexico to Venezuela and Trinidad & Tobago, but not specifically from Costa Rica
(Bacon 1956, Hynes 1971, Roback and Nieser 1974). Reported here are the first records from Costa Rica.
Material examined. COSTA RICA: Alajuela—San Carlos, San Carlos River Basin, TEC Project, station 2,
28.VII.1999: 3 apterous males, 1 macropterous male, 2 apterous females, 2 macropterous females; station 3,
25.VIII.1999: 2 apterous males, 2 apterous females, 1 nymph. San Carlos, Arenal River below Vado, 24.II.2011, (F.
Reyes & D. Vázquez): 3 apterous males. Guanacaste—Palo Verde, Lomas Barbudal Corridor, 15.I.2007, (B.
Pacheco): 1 apterous male. Bagaces, Lomas Barbudal, river, 300 m a.s.l., 4.X.1998, (P. Ortiz): 5 apterous males, 1
macropterous male, 1 apterous female. Limón—Valle de La Estrella, Estrella River, 1.5 km downstream levee,
12.VII.2012, (B. Pacheco & F. Reyes): 2 apterous males, 4 apterous females, 1 macropterous female, 6 nymphs.
Limón, Portete, stream, under 50 m a.s.l., 22.IV.1995, (L. Vik): 3 apterous females, 11 nymphs. Limón, Japdeva
Channel, 02.V.2013, (B. Pacheco & F. Garita): 2 apterous males. Puntarenas—Osa, Piro River, 8o24.117’ N / 83o
20.353’ W, 32 m a.s.l., 23.V.2009, (B. Pacheco): 1 macropterous male, 1 apterous female. Osa, Rincón, Rincón
River, 15 m a.s.l., 11.X.2005, (J. Bermúdez): 2 apterous males. Osa, Osa Peninsula, Rincón River, bridge before
mouth, 20 m a.s.l., 27.IX.1996, (M. Springer): 1 apterous male.

Rhagovelia crassipes Champion, 1898

(Fig. 30)

Known distribution in Costa Rica. Alajuela, Cartago, Guanacaste, Puntarenas, and San José (Torre-Bueno 1906,
Polhemus 1997). Reported here is the first record from Heredia Province.
Material examined. COSTA RICA: Alajuela—Palmares, Aguacate Stream, 10o1.208’ N / 84o27.825 W,
12.IV.2009, (B. Pacheco): 2 macropterous females. Palmares, La Catarata Stream, Rincón de Zaragoza, 1100 m
a.s.l., 14.VIII.2005, (B. Pacheco): 1 apterous male, 1 apterous female. Zaragoza, waterfall, 1200 m a.s.l., 12.I.2005,
(F. Ruíz): 2 apterous males, 1 apterous female, 1 macropterous female. San Ramón, REBAMB, San Lorencito
River, 900 m a.s.l., 24.IX.2005, (M. Alfaro): 1 apterous male. San Ramón, Quebradillas, El Salto Stream, 1100 m
a.s.l., 14.X.2005, (A. Castillo): 1 apterous female. Heredia—Santa Bárbara, Cachazas Stream, 1100 m a.s.l.,
16.X.1998, (G. Chaverri): 1 apterous male, 1 apterous female. Braulio Carrillo, stream 200 m behind La Fonda
Restaurant, 1600 m a.s.l., 30.XI.1996, (N. Franz): 1 apterous male, 1 apterous female. San José—María Aguilar
River, 7.I.2003: 1 apterous female; 7.IX.2003, (A. Y. Jiménez): 1 apterous male, 3 apterous females, 1
macropterous female. Moravia, Zurquí, Yerbabuena Stream, 1600 m a.s.l., III.1995, (C. Flores): 3 apterous males,
3 apterous females, 3 nymphs; 2.IV.1995, (J. Coronado): 5 apterous males; 1500 m a.s.l., IV.1995, (P. Hanson): 5
apterous males, 3 apterous females.

Rhagovelia elegans Uhler, 1894

(Fig. 32)

Known distribution in Costa Rica. Puntarenas and San José (Polhemus 1997). Reported here are the first records
from Alajuela and Guanacaste provinces.
Material examined. COSTA RICA: Alajuela—San Carlos, Aguas Zarcas, Altamira, Mr. Ash Stream, 100 m
a.s.l., 2010, (B. Pacheco): 1 apterous female. Guanacaste—Cañas, Palo Verde National Park, ponds on sides of
trail to Tárcoles River, 28–29.X.2005, (P. Calderón): 2 apterous males, 2 apterous females. Nicoya, Barra Honda
National Park, Ojo de Agua el Anís, 80 m a.s.l., 11.II.2005, (F. Ruíz-Pacheco): 1 macropterous female; Barra
Honda National Park, La Cascada, 110 m a.s.l., 8.I.2005, (F. Ruíz-Pacheco): 1 macropterous female, 1 nymph.
Puntarenas—Parrita, Parritón Stream, site 5, 9o38.980’ N / 84o15.986 W, 11.V.2009, (B. Pacheco): 2 macropterous
males. Golfito, Cañaza Stream, under waterfall, I.1997, (A. Villalobos): 6 macropterous males, 7 macropterous
females. Santa Elena, Berros Stream, 1200 m a.s.l., IV.2001, (A. Burgin): 1 macropterous female.

TWO NEW SPECIES OF RHAGOVELIA Zootaxa 3980 (4) © 2015 Magnolia Press · 491
Rhagovelia femoralis Champion, 1898
(Fig. 33)

Known distribution in Costa Rica. Alajuela, Heredia, and Puntarenas (Polhemus 1997). Reported here are the
first records from Cartago and San José provinces.
Material examined. COSTA RICA: Alajuela—San Ramón, Alberto Manuel Brenes Biological Reserve, 700
m a.s.l., 23.III.2007, (B. Pacheco & M. Zumbado): 2 apterous males, 1 macropterous male, 2 nymphs; Alberto
Manuel Brenes Biological Reserve, San Lorencito River, 10o13’6.8” N / 84o35’48.9” W, 876 m a.s.l., 17.X.2009,
(B. Pacheco): 3 apterous males, 1 apterous female; San Lorencito River, in front of station, 860 m a.s.l., 16.II.1994,
(M. Springer): 1 apterous male, 3 apterous females, 5 nymphs; San Lorencito River, 950 m a.s.l., 25.IX.2005, (A.
Castillo): 1 apterous female. San Ramón, REBAMB, 18.VIII.2003: 1 apterous male, 2 apterous females. San
Ramón, REBAMB, San Lorencito River, 900 m a.s.l., 4.X.2003, (C. Boucher): 1 apterous male, 1 apterous female.
San Ramón, San Ramón Forest Reserve, 850 m a.s.l., 25.III.1995, (J. Ludwig & R. L. Rodríguez): 2 apterous
males, 3 apterous females. San Ramón, San Ramón Reserve, San Lorencito River, 23.XI.1996, (M. Springer): 10
apterous males, 10 apterous females, 22 nymphs. San Ramón, Piedades Sur, Yayo Stream, 700 m a.s.l., 21.IX.2003,
(R. Lara): 1 apterous male, 1 apterous female. San Ramón, San Lorenzo River, 1500 m a.s.l., 5.IX.2003, (F. Ruíz-
Pacheco): 1 apterous male, 1 nymph. Palmares, La Catarata Stream, Rincón de Zaragoza, 1100 m a.s.l.,
14.VIII.2005, (B. Pacheco): 1 apterous female. San Isidro, Concepción, Tranqueras River, 1600 m a.s.l.,
26.XI.2000, (E. Bermúdez): 2 apterous males. Cartago—La Unión, Tres Ríos, Cerro La Carpintera, Istarú, La
Chirraca Stream, 1555 m a.s.l., 6.IX.1999, (S. Jenes): 4 apterous males, 4 apterous females. Turrialba, Jícotea,
Jícotea River, 1000 m a.s.l., 17.VII.1991, (M. Springer): 1 apterous male, 1 macropterous female; 09.VI.1991, (M.
Springer): 1 apterous male, 2 macropterous males, 1 apterous female; Jícotea River, 850 m a.s.l., 16.VII.1991 (M.
Springer): 1 apterous male, 1 apterous female. Paraíso, Birrisito, Parruas River Stream, 1350 m a.s.l., 15.X.1996,
(C. Guillén): 2 apterous males. Heredia—Sarapiquí, Angel River, 1400 m a.s.l., 04.X.1998, (T. Mora): 3 apterous
males, 3 apterous females, 5 nymphs. Sarapiquí, Cariblanco, stream tributary to Sarapiquí River, 900 m a.s.l.,
25.VIII.2005, (B. Pacheco): 2 apterous males, 1 macropterous male, 1 apterous female. San Isidro, Lajas River,
1300 m a.s.l., III.1995, (V. Solano): 1 apterous male. Zurquí, Yerbabuena Stream, 1500 m a.s.l., IV.1995, (P.
Hanson): 1 apterous male. Puntarenas—Parrita, Damitas River, 9o33.559’ N / 84o11.480’ W, 14.VI.2009, (B.
Pacheco): 1 teneral apterous male, 2 macropterous males, 2 macropterous females, 1 nymph. Golfito, Cañaza
Stream, under waterfall, under 50 m a.s.l., I.1997, (A. Villalobos): 1 apterous male, 1 apterous female, 1
macropterous female; Cañaza Stream, waterfall, 10 m a.s.l., IV–V.1999, (T. Hermanson): 1 teneral male. Golfito,
El Naranjal Stream, 10 m a.s.l., 20–21.I.2000, (M. Springer): 1 apterous female, 1 nymph. Buenos Aires, Longo
Mai, Sonador River, 1050 m a.s.l., 23.VIII.1998, (A. Nunn): 1 apterous male. Golfito, La Gamba Station, Chorro
Stream, under 100 m a.s.l., (19.I.2011), (R. Naumann): 1 apterous male, 1 apterous female, 1 nymph. Puntarenas,
San Luis, Monteverde, Alondra Stream, 1200 m a.s.l., 23.VI.1995: 10 apterous males, 2 macropterous males, 6
apterous females, 2 macropterous female, 17 nymphs. Coto Brus, Las Cruces, Jaba River (Matadero), 10.X.1999,
(M. Springer): 1 apterous male. Coto Brus, Las Cruces, Jaba River, 1100 m a.s.l., (M. Springer): 1 apterous male, 1
macropterous male; 1200 m a.s.l., (II.1998): 2 apterous males, 2 apterous females, 1 female with broken wings, 6
nymphs. San José—Acosta, Palmichal, Tabarcia River and Negro River, 25.II.2009, (B. Pacheco): 2 apterous
females. Acosta, San Ignacio, Chirraca Waterfalls, 1100 m a.s.l., 9.X.2005, (E. Arroyo): 1 apterous male. Moravia,
Pará-Blanco River, 10o1.610 N / 84o0.665’ W, 1418 m a.s.l., 1.II.2009, (B. Pacheco): 2 apterous females, 4 nymphs.
Mora, Finca El Rodeo, lake, 04.XI.2000, (D. Vásquez): 1 apterous female. Santa Ana, La Cruz Stream, 1800 m
a.s.l., 1.XI.2003, (C. Boucher): 1 apterous male, 1 apterous female.

Rhagovelia formosa Bacon, 1956

(Fig. 31)

Known distribution in Costa Rica. Cartago, Puntarenas, and San José (Polhemus 1997).
Material examined. COSTA RICA: Puntarenas—Lepanto, Blanco River, Paso Tres Ríos, 350 m a.s.l.,
23.I.1998: 5 macropterous males, 6 macropterous females.

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Rhagovelia scabra Bacon, 1956
(Fig. 30)

Known distribution in Costa Rica. Alajuela, Cartago, Heredia, Puntarenas, and San José (Bacon 1956; Wilson et
al. 1978; Polhemus 1997).
Material examined. COSTA RICA: Alajuela—San Carlos, Quesada, San Vicente, Peje River, 1500 m a.s.l.,
14.VIII.2005, (P. Benavides): 1 apterous male. San Ramón, REBAMB, 18.VIII.2003: 2 apterous males; REBAMB,
1100 m a.s.l., 18.X.2003, (F. Ruíz-Pacheco): 1 macropterous male. Cartago—Tapantí, Orosi River, 23.X.2005,
(M. Werner): 3 apterous males, 2 apterous females. Valle del Guarco, Palo Verde del Guarco, 10.IV.2009, (A.
Bourg): 2 apterous males, 3 apterous females. La Unión, Tiribí River, 1780 m a.s.l., 20–23.VIII.1998, (R. Mora): 3
apterous males, 1 macropterous male, 3 apterous females, 1 macropterous female, 3 nymphs. La Unión, Tres Ríos,
7.IX.2003: 3 apterous males, 1 macropterous male, 2 apterous females. Orosi, tributary of Grande de Orosi River,
between Purisil and Tapantí, 20.IX.1998, (F. Mejía): 1 macropterous male. Orosi, Tapantí, Grande de Orosi River,
Oropendula Trail, 1240 m a.s.l., 13.XI.1999, (M. Springer): 10 apterous males, 1 macropterous male, 3 apterous
females, 3 macropterous females. Orosi, Tapantí Stream: 1 apterous male, 3 apterous females. Oreamuno, Cerro La
Carpintera, Istarú, La Chirraca Stream, 1555 m a.s.l., 6.IX.1999, (S. Jenes): 10 apterous males, 9 apterous females.
Heredia—San Rafael, Los Turales River, 8.II.1997, (C. Ruiz & P. Cambronero): 3 apterous males, 1 macropterous
male, 1 apterous female. San Rafael, Cerro Chompipe, Riachuelo, 200 m a.s.l., 2.IX.2000: 2 apterous males. San
Rafael, La Bermúdez Stream, 8.II.1997, (C. Ruiz & P. Cambronero): 2 apterous males, 2 macropterous males, 1
macropterous female, 1 nymph. San Isidro, B. Santa Cecilia, Lajas River, 1700 m a.s.l., 6.XII.2003, (J. Sibaja): 1
apterous male, 1 apterous female, 1 nymph. San Isidro, Concepción, Tranqueras River, 26.XI.2000, (E. Bermúdez):
6 apterous males, 6 apterous females. Tibás River, 1600 m a.s.l., 1.XII.2000: 2 apterous males, 1 apterous female, 1
macropterous male, 5 nymphs. La Amistad Stream, 10o4’ N / 84o7’ W, between Porrosati and Sacramento, 1900 m
a.s.l., V–VI.1999, (T. Klug): several apterous males and females. Puntarenas—Pérez Zeledón, Herradura, Rivas,
Blanco River, 9o30.008 N / 83o36.783 W, (D. Zamora): 2 apterous males. Puntarenas, Monteverde, 1400 m a.s.l.,
4.2001, (A. Burgin): 2 apterous males, 1 apterous female, 4 nymphs. Puntarenas, Monteverde, lake in front of La
Cascada, 1400 m a.s.l., 1.III.1998, (M. Springer): 1 apterous male. San José—Cerro de La Muerte, margins of
river, 9.73888o N / 83.98446o W, 2009, (B. Pacheco): 1 apterous female. Cerro de La Muerte, spout by Soda Los
Traileros, 16.IX.2007, (B. Pacheco): 1 apterous male, 3 apterous females, 1 nymph. María Aguillar River, 7.I.2003:
1 apterous male; 7.IX.2003: 1 apterous female. Escazú, San Antonio, Agres River, 1300 m a.s.l., 04.VI.1995, (L.
Vik & I. Coronado): 8 apterous males, 1 macropterous male, 8 apterous females, 2 macropterous females.
Coronado, Los Juncos, Grande Stream, 1600 m a.s.l., 20.IX.2005, (E. Boza): 1 apterous female. Coronado,
Cascajal, Derrumbe Stream, approximately 1600 m a.s.l., 22.X.2005, (M. Losilla): 1 apterous male, 1 apterous
female. Montes de Oca, San Rafael, Mansiones, Torres River, 10.XII.2000, (S. Carvajal): 3 apterous males, 3
apterous female, 1 teneral female, 1 nymph.

Rhagovelia solida Bacon, 1956

(Fig. 30)

Known distribution in Costa Rica. Heredia and Puntarenas (Polhemus 1997). Reported here is the first record
from San José Province.
Material examined. COSTA RICA: Heredia—La Amistad Stream, 10o4’ N / 84o7’ W, between Porrosati and
Sacramento, 1900 m a.s.l., III–IV.1999, (T. Klug): 1 apterous male, 5 apterous females, 7 nymphs. Bosque de la
Hoja Stream, 1400 m a.s.l., 5.III.1996, (C. Ugalde): 7 apterous males, 5 apterous females, 1 nymph. San José—
Coronado, Juncos, Cascajal River, 1500 m a.s.l., 8.X.2005, (R. Sánchez): 1 apterous female, 1 nymph.

Rhagovelia spinigera Champion, 1898

(Fig. 31)

Known distribution in Costa Rica. San José, Río Virilla [unknown province], and Pavas [unknown province]

TWO NEW SPECIES OF RHAGOVELIA Zootaxa 3980 (4) © 2015 Magnolia Press · 493
(Torre-Bueno 1906, Bacon 1956, Polhemus 1997). Reported here are the first records from Alajuela, Cartago, and
Heredia provinces.
Material examined. Alajuela—San Ramón, Alto Zamora, 1200 m a.s.l., IX.2005, (R. Gómez): 1
macropterous male, 1 apterous female. Cartago—Orosi, 22.XI.2008, (B. Pacheco): 7 apterous males, 1
macropterous male, 2 apterous females. Orosi, arm of Grande de Orosi River, between Purisil and Tapantí,
20.IX.1998, (F. Mejía): 1 apterous male. Orosi, Porisi River, strong current, 20.IX.1998: 1 apterous female, 3
nymphs. Orosi, waterfall under bridge between Orosi and Paraíso, 20.IX.1998, (F. Mejía): 2 apterous males, 2
apterous females, 1 nymph. Orosi, polluted waterfall, 20.IX.1998, (C. Obardo): 8 apterous males, 2 macropterous
males, 8 apterous females. Heredia—San Isidro, Santa Elena, Yerbabuena River, 390 m a.s.l., 6.IX.1998, (A.
León): 1 apterous female.

Rhagovelia spinosa Gould, 1931

(Fig. 28)

Previously recorded from Honduras to Peru, but not specifically from Costa Rica (Gould 1931, Bacon 1956, Drake
and Hussey 1957, Padilla-Gil 2011). Reported here are the first records from Costa Rica.
Material examined. Alajuela—San Carlos, San Carlos River Basin, TEC Project, VII–IX.1999: 1 apterous
male, 1 macropterous female. Limón—Valle de la Estrella, Estrella River, VII.2012, (B. Pacheco & F. Reyes): 3
apterous male, 3 apterous females. Puntarenas—Cuenca del Térraba, Diquís Project, ICE, General River X Peje
River, 23.III.2006, (R. Lara): 14 apterous males, 11 apterous females.

Rhagovelia tantilla Drake & Harris, 1933

(Fig. 28)

Previously recorded from Belize to Peru, but not specifically from Costa Rica (Drake and Harris 1933, Bacon
1956). Reported here are the first records from Costa Rica.
Material examined. COSTA RICA: Alajuela—San Carlos, Aguas Zarcas, Altamira, Mr. Ash Stream, 100 m
a.s.l., 2010, (B. Pacheco): 1 apterous female. San Ramón, San Lorencito River, 10.X.1998: 1 macropterous female.
Guanacaste—Hojancha, Nosara River, 850 m a.s.l., 14.IX.2003, (J. A. Vindas): 1 male with broken wings. ACG,
Góngora River, under bridge, on way to Cerro Cacao Reserve, 900 m a.s.l., 5.X.1998, (M. Springer): 1 apterous
male, 1 apterous female. Heredia—Sarapiquí, Tirimbina Reserve, El Uno Stream, 3.X.2005, (M. Werner): 6
apterous males, 1 apterous female. Sarapiquí, La Selva Biological Station, Trail SSE, Sábalo Stream, 50 m a.s.l.,
VI.1997: 1 apterous male. Limón—Braulio Carrillo National Park, Botarrama Stream, González Stream Sector,
29.I.2006: 6 apterous males, 2 apterous females; Braulio Carrillo National Park, Botarrama Stream, 9.X.2005: 1
apterous male, 1 apterous female. Matina, Brazo Seco Stream, 10.I.2012, (B. Pacheco & F. Reyes): 1 apterous
male. Pococí, Guápiles, Santa Clara, 400 m a.s.l., 14.IX.2003, (S. Amador): 1 apterous male. Pococí, Guápiles,
Guápiles River, 550 m a.s.l., 5.VIII.1998, (L. Q. Fernãndez): 1 apterous male. Puntarenas—Osa, Piro River,
8o24.117’ N / 83o20.353 W, 23.V.2009, (B. Pacheco): 2 apterous females. Osa, Osa Peninsula, tributary to Sábalo
Stream, bridge on road to Puerto Jimenez, under 20 m a.s.l., 27.IX.1996, (M. Springer): 2 apterous females; Osa
Peninsula, Carbonera River, 250 m a.s.l., 23.XII.1996, (M. Springer): 1 apterous male, 3 nymphs. Zona Sur,
Interamericana, Coto River, by the bridge, under 50 m a.s.l., X.1997, (G. Umaña): 1 apterous male, 3 nymphs.
Golfito, Cañaza Stream, 10 m a.s.l., 16.I.1998, (M. Springer): 3 apterous males, 6 apterous females;
Interamericana, Esquinas River, bridge, under 50 m a.s.l., 19.VII.1996, (G. Umaña): 1 apterous male. Buenos
Aires, Longo Mai, San Juan Rive, 550 m a.s.l., 24.IX.1998, (A. Nunn): 2 apterous females. Puntarenas, Chomes,
Lagarto River, 4.XI.2000, (A. Loría): 1 apterous male, 4 apterous females, 1 nymph. San José—Coronado,
Cascajal, Cascajal Stream, 750 m a.s.l., 28.IX.2003, (S. Amador): 1 apterous female.

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FIGURES 28 & 29. Geographical distribution of the collecting localities of Rhagovelia in Costa Rica. 28. Rhagovelia
angustipes, R. danpolhemi sp. nov., R. calopa, R. spinosa, R. tenuipes, and R. tantilla. 29. R. springerae sp. nov.

TWO NEW SPECIES OF RHAGOVELIA Zootaxa 3980 (4) © 2015 Magnolia Press · 495
FIGURES 30 & 31. Geographical distribution of the collecting localities of Rhagovelia in Costa Rica. 30. Rhagovelia
acuminata, R. armata, R. scabra, R. solida, and R. crassipes. 31. R. formosa, R. spinigera, and R. bisignata.

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FIGURES 32 & 33. Geographical distribution of the collecting localities of Rhagovelia in Costa Rica. 32. Rhagovelia
uncinata and R. elegans. 33. R. femoralis.

TWO NEW SPECIES OF RHAGOVELIA Zootaxa 3980 (4) © 2015 Magnolia Press · 497
Rhagovelia tenuipes Champion, 1898
(Fig. 28)

Known distribution in Costa Rica. San José (Hungerford 1939, Bacon 1956). Reported here are the first records
from Alajuela, Guanacaste, Heredia, Limón, and Puntarenas provinces.
Material examined. COSTA RICA: Alajuela—San Carlos, Florencia, La Vieja River, powerhouse of
Hydropower Station Chocosuela II, 325 m a.s.l., 15.VI.2002, (R. Acosta): 1 male with broken wings, 1 nymph. San
Carlos, San Carlos River Basin, TEC Project, site 1, VII–IX.1999: 1 apterous male, 1 macropterous male, 1
apterous female; site 7, 18.VIII.1999: 1 apterous male; site 6, 21.VII.1999: 1 macropterous male, 1 macropterous
female. San Carlos, Cutris, Crucitas, Relaves, 10.86662o N / 84.34776o W, X.2008, (B. Pacheco): 1 apterous male.
San Carlos, Celeste River, 10o43.843’ N / 84o53.270’ W, 106 m a.s.l., 23.IV.2008, (B. Pacheco): 1 apterous male, 1
apterous female. San Carlos, Aguas Zarcas, Altamira, Mr. Ash Stream, 100 m a.s.l., 2010, (B. Pacheco): 2 apterous
females. San Carlos, San Pedro (La Tigra), small tributary of Peñas Blancas River, 150 m a.s.l., 20.X.2001, (M.
Springer & C. Guillén): 1 apterous male. San Carlos, Mojahuevos Stream, 100 m a.s.l., 30.III.2002, (K. S.
Rodríguez): 7 apterous males, 8 apterous females. San Ramón, San Ramón Reserve, San Lorencito River, 860 m
a.s.l., 16.II.1994, (M. Springer): 1 apterous female. San Ramón, Alberto Manuel Brenes Mora Reserve, San
Lorencito River, 950 m a.s.l., 25.X.2003, (J. A. Vindas): 2 apterous females; 25.IX.2005, (A. Castillo): 1 male with
broken wings. San Ramón, San Lorencito River, 10.X.1998: 1 apterous female. San Ramón, San Ramón Forest
Reserve, 850 m a.s.l, 25.III.1995, (J. Ludwig & R. L. Rodríguez): 5 apterous males, 3 apterous females. San
Ramón, Las Musas River, 850 m a.s.l, 31.VIII.2003, (A. Y. Jiménez): 1 apterous male, 2 apterous females, 1
nymph. San Ramón, Piedades Sur, Yayo Stream, 700 m a.s.l., 21.IX.2003, (R. Lara): 1 apterous male, 1 apterous
female. Upala, Canalete, Achiote River, desfogue, VIII.2010, (F. Reyes & P. Gutiérrez): 4 apterous females. Upala,
Canalete, Zapote River, site 6, desfogue, (B. Pacheco): 1 apterous male. Guanacaste—Palo Verde, Lomas
Barbudal Corridor, Barbudal Stream, 15.I.2007, (B. Pacheco): 1 apterous male, 1 apterous female. ACG, Góngora
River, under bridge, on way to Cerro Cacao Reserve, 900 m a.s.l., 5.X.1998, (M. Springer): 3 apterous females.
Cañas, Corobicí River, 30.VII.1994, (C. Angulo): 1 apterous male, 1 apterous female. Cañas, site 6, La Palma
Stream, EsIA south channel, 54 m a.s.l., VII.2010, (B. Pacheco & D. Vásquez): 1 apterous female; site 7, La
Soledad Stream, EsIA south channel, 51 m a.s.l., VII.2010, (B. Pacheco & D. Vásquez): 1 apterous female, 1
macropterous female. Abangares, Las Juntas, Ecomuseo, Aguas Claras River, 500 m a.s.l., 17.IX.2005, (E.
Arroyo): 1 male with broken wings, 2 apterous females, 5 nymphs. Tilarán, Rugama Stream, 650 m a.s.l.,
09.V.2011, (B. Pacheco, D. Vásquez & J. Bermúdez): 1 apterous male, 3 apterous females. Heredia—Braulio
Carrillo National Park, 20.X.2007, (S. Zamora): 1 male with broken wings, 1 apterous female; Braulio Carrillo
National Park, Molenete Stream, cableway, 600 m a.s.l., 10.XI.1996, (E. Estrada): 1 apterous male, 1 apterous
female. Sarapiquí, La Tirimbina Reserve, Sarapiquí River, 27.I.2006: 1 apterous female, 1 nymph; 3.X.2005, (M.
Werner): 2 apterous males; Tirimbina Reserve, El Uno Stream, 3.X.2005, (M. Werner): 1 apterous male, 1 apterous
female. Sarapiquí, Selva Verde Lodge, Sarapiquí River, 100 m a.s.l., 30.IV.2000, (M. Springer): 1 apterous male, 3
apterous females, 1 macropterous female. Sarapiquí, Poza Azúl River, site 3, 275 m a.s.l., III.1998, (P. Paaby): 1
apterous female, 1 macropterous male. Sarapiquí, La Virgen, tributary to Poza Azúl River, 250 m a.s.l., III.1998, (P.
Paaby): 1 apterous male, 2 nymphs. Horquetas, above ICE Project, stream behind Horquetas gas station,
19.VII.2012, (B. Pacheco): 1 apterous male, 2 apterous females, 2 nymphs. Limón—Peje River, Reventazón
Project, site 10, XI.2011, (B. Pacheco & F. Reyes): 2 apterous males, 6 apterous females. Valle de La Estrella,
Estrella River, VII.2012, (B. Pacheco & F. Reyes): 1 apterous male. Braulio Carrillo National Park, Botarrama
Stream, González Stream Sector, 29.I.2006: 7 apterous males, 11 apterous females; Botarrama Stream, 9.X.2005: 4
apterous males, 5 apterous females. Matina, Brazo Seco Stream, 10.I.2012, (B. Pacheco & F. Reyes): 1 apterous
female. Pococí, Guacimito River, 550 m a.s.l., 19.XI.2000, (L. Rodríguez): 1 apterous male, 1 male with broken
wings, 12 apterous females, 1 female with broken wings, 9 nymphs. Pococí, Guápiles, Costa Rica River, 230 m
a.s.l., 14.XI.2005, (E. Arroyo & G. Vargas): 1 apterous male, 2 apterous females; 14.X.2005, (T. Cotta): 1 female
with broken wings; Costa Rica River, approximately 200 m a.s.l., (P. Calderón): 1 apterous male, 1 apterous
female; Costa Rica River, under bridge, 14.XI.2005, (C. Lizana): 1 female with broken wings. Pococí, La Earth,
Novillos River, 40 m a.s.l., 22.V.1996, (E. Martínez): 1 apterous female, 1 macropterous female, 7 nymphs. La
Argentina, Finca Los Manantiales, stream, 570 m a.s.l., 21.VIII.2005, (M. Montero-Guzmán): 1 macropterous
female. Guápiles, Chirripó River, site 1, above CDP, 07.IV.2011, (B. Pacheco): 3 apterous males, 1 macropterous
male, 2 apterous females; Chirripó River, inside CDP: 2 apterous females. Limón, unnamed stream, tributary to
Limoncito River, (B. Pacheco & F. Garita): 2 apterous females. Puntarenas—Osa, Banegas River, 8o41.485’ N /

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83o32.211 W, 31.I.2008, (B. Pacheco): 1 apterous male, 1 macropterous female. Osa, Puerto Jiménez, Tigre River,
25.II.2010, (B. Pacheco): 8 apterous males, 2 apterous females, 2 nymphs. Osa, Corcovado National Park,
8o28.703’ N / 83o35.447 W, XI.2009 (B. Pacheco): 1 apterous male. Osa, Piedras Blancas River, 2.II.2008, (B.
Pacheco): 2 apterous males. Osa, Osa Peninsula, Conte River, bridge, on way to Puerto Jiménez, under 20 m a.s.l.,
27.IX.1996, (M. Springer): 2 apterous males, 3 nymphs; Osa Peninsula, tributary to Sábalo River, under 20 m a.s.l.,
27.IX.1996, (M. Springer): 1 apterous male; Osa Peninsula, Barrigones River, by the bridge, under 50 m a.s.l.,
20.VII.1996, (G. Umaña): 1 macropterous male, 1 nymph; Osa Peninsula, Tigre River, under 50 m a.s.l.,
20.VII.1996, (G. Umaña): 1 apterous male; Osa Peninsula, Agujas River, by the bridge, under 50 m a.s.l.,
26.IV.1996, (G. Umaña): 1 apterous female, 4 nymphs; Osa Peninsula, Corcovado National Park, Claro River, near
Sirena, 0 m a.s.l., 24.IV.1999, (L. Stauffer): 5 apterous males, 1 macropterous male, 5 apterous females. Osa, Brujo
River, 0 m a.s.l., 15.IX.2005, (T. Cotta): 1 female with broken wings. Osa, Ramón River, Bio Terra, IV.2011, (D.
Zúñiga): 1 apterous female, 1 macropterous female; (B. Pacheco): 1 apterous female. Cuenca del Térraba, Térraba
River, IV.2000: 1 apterous male, 1 male with broken wings, 1 apterous female. Cuenca Del Térraba, Diquís
Hydropower Project, Convento River, 25.II.2005, (R. Lara): 4 apterous males, 10 apterous females; Diquís
Hydropower Project, Brujo River, 1.III.2006, (R. Lara): 2 apterous males, 1 macropterous female; Diquís
Hydropower Project, ICE, General River X Peje River, 23.III.2006, (R. Lara): 1 apterous male, 3 apterous females.
Diquís Hydropower Project, Unión River, 14.III.2006, (R. Lara): 2 apterous males, 2 males with broken wings, 2
apterous females, 1 female with broken wings. Diquís Hydropower Project, Volcán River, 13.II.2006, (R. Lara): 3
apterous females. Parrita, Bijagual, Pirrís River, 8o38.890 N / 84o15.986 W, 14.V.2009, (B. Pacheco): 1 apterous
male, 1 macropterous male, 6 apterous females, 1 macropterous female. Punta Morales, Lagarto River, 10o6.532’ /
84o55.132’ W, 38 m a.s.l., 7.VI.2009, (B. Pacheco): 2 apterous males, 1 apterous female. Buenos Aires, Longo Mai,
Convento River, 550 m a.s.l., 23.V.1998, (A. Nunn): 1 apterous male; 23.VII.1998, (A. Nunn): 1 apterous female;
Longo Mai, San Juan River, 550 m a.s.l., 24.IX.1998, (A. Nunn): 1 apterous male, 3 apterous females. Golfito,
Cañaza Stream, under waterfall, under 50 m a.s.l., I.1997, (A. Villalobos): 1 apterous female, 1 female with broken
wings, 1 macropterous female. Golfito, La Gamba Stream, 10 m a.s.l., 15.I.1998, (M. Springer): 1 apterous male.
Montes de Oro, Miramar, Ciruelas River, 500 m a.s.l, 10.X.2005, (W. Solis): 1 apterous female. Zona Sur,
Interamericana, Coto River, by the bridge, under 50 m a.s.l, 25.IV.1997, (G. Umaña): 1 apterous female. Coto Brus,
Las Cruces, Jaba River (Matadero), 10.X.1999, (M. Springer): 1 apterous male. Coto Brus, San Vito, Las Cruces,
Jaba River, 1100 m a.s.l., (M. Springer): 1 macropterous female. San José—Pérez Zeledón, San Isidro, stream, 700
m a.s.l., 15.IX.2005, (R. Lara): 2 apterous females, 3 nymphs.

Rhagovelia uncinata Champion, 1898

(Fig. 32)

Known distribution in Costa Rica. Puntarenas and San José (Hungerford 1939, Polhemus 1997). Reported here
are the first records from Alajuela, Guanacaste, Heredia, and Limón provinces.
Material examined. COSTA RICA: Alajuela—Rugama Stream, Hydropower Project Cote, 19.V.2008, (B.
Pacheco): 1 macropterous female. Upala, Hydropower Project Canalete, 10o46.495’ N / 85o2.516’ W, 245–265 m
a.s.l., 24.XII.2008, (B. Pacheco): 1 apterous male, 2 apterous females. Upala, site 4, Tuberíunbre, Bijagua River,
canalete, VIII.2010, (F. Reyes & P. Gutiérrez): 1 macropterous female. Guanacaste—Nicoya, Ojo de Agua el
Bejuco, 95 m a.s.l., 10.I.2005, (F. Ruíz-Pacheco): 1 apterous male, 1 apterous female. Nicoya, Barra Honda
National Park, La Caña River, 225 m a.s.l., 15.I.2005, (F. Ruíz-Pacheco): 1 apterous male. Heredia—Sarapiquí,
Sarapiquí River, under bridge, La Tirimbina Reserve, 3.VI.2008, (B. Pacheco): 1 apterous male. Sarapiquí,
Tirimbina Station, Sarapiquí River, 3.X.2005, (M. Werner): 1 apterous female. Sarapiquí, Puerto Viejo,
Organization for Tropical Studies, La Selva Station, stream, 50 m a.s.l., 23.IX.1996, (E. Martínez): 1 apterous
female; La Selva Station, CCL Trail, El Salto Stream, 50 m a.s.l., VI.1996: 1 apterous male, 2 apterous females, 7
nymphs; VI.1997: 1 apterous male, 1 apterous female, 1 nymph. Limón—La Argentina, Finca Los Manantiales,
stream, 570 m a.s.l., 21.VIII.2005, (M. Montero-Guzmán): 1 apterous female. Puntarenas—Coto Brus,
Organization for Tropical Studies, Las Cruces Station, 22.I.2010, (M. Springer): 1 macropterous male; Las Cruces,
Jaba River (Matadero), 10.X.1999, (M. Springer): 1 apterous male, 1 apterous female; Jaba River, 1100 m a.s.l.,
(M. Springer): 1 macropterous male, 1 macropterous female. Osa, Corcovado National Park, 8o28.703’ N /
83o35.447’ W, XI.2009, (B. Pacheco): 1 apterous female. Osa, Peninsula de Osa, pond in forest, 5 m a.s.l.,

TWO NEW SPECIES OF RHAGOVELIA Zootaxa 3980 (4) © 2015 Magnolia Press · 499
28.X.2000, (E. Bermúdez): 1 apterous male. Parrita, Parritón Stream, 9o38.980’ N / 84o15.986’ W, 11.V.2009, (B.
Pacheco): 9 macropterous males, 2 macropterous females. Golfito, Cañaza Stream, 10 m a.s.l., I.1997, (A.
Villalobos): 2 macropterous males, 2 macropterous females; 16.I.1998, (M. Springer): 1 apterous male, 1
macropterous male, 9 nymphs; IV–V.1999, (T. Hermanson): 1 apterous female; Cañaza Stream, bridge leaving
Golfito, 0 m a.s.l., I.1997, (A. Villalobos): 1 macropterous male, 1 apterous female. Golfito, La Gamba, Esquinas
Waterfall Trail, second and third waterfalls, 19.II.2011, (D. Takiya): 2 apterous males, 6 apterous females, 2
nymphs. Golfito, La Gamba Stream, 2 km from Barrio Ureña, primary forest, 120 m a.s.l., I.1997: 2 apterous
males, 1 macropterous male, 5 apterous females. Finca La Esperanza, near Esquinas River, clear stream, 100 m
a.s.l., 9.VI.1995, (L. Vik): 2 apterous males, 2 nymphs. Puntarenas, Montezuma, waterfall, 0 m a.s.l., IV.1995, (C.
Nolting): 1 macropterous male, 1 apterous female. Puntarenas, Lepanto, Blanco River, Paso Tres Ríos, 350 m a.s.l.,
23.I.1998: 1 macropterous male. Puntarenas, Monteverde, lake in front of La Cascada, 1400 m a.s.l. 1.III.1998, (M.
Springer): 2 apterous females. San José—Acosta, Palmichal, Tabarcia River and Negro River, 1200 m a.s.l.,
25.II.2009, (B. Pacheco): 1 macropterous female. Acosta, San Ignacio, Chirraca Waterfall, 1100 m a.s.l., 9.X.2005,
(E. Arroyo): 2 apterous males. Mora, Finca El Rodeo, lake, 4.XI.2000, (D. Vásquez): 1 apterous male.

Acknowledgements

We thank Fátima Reyes Morales for her valuable help with the laboratory work and Darha Solano for providing
Figures 2–3. Collecting permits were provided by the National System of Conservation Areas (SINAC) and Raquel
Romero helped with the elaboration of the distribution maps. The manuscript of this article benefited from the
useful comments of Dr. M.B. Malipatil, Dr. Robert W. Sites (University of Missouri), and an anonymous reviewer.
This publication was made possible due to the contribution of many researchers and students who deposited their
specimens over the last two decades at the Museo de Zoología of the Universidad de Costa Rica.

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