Perkins 2020 Escarabajos Buceadores de Tambopata 10.11646@zootaxa.4868.1.4

Zootaxa 4868 (1): 061–089 ISSN 1175-5326 (print edition)
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Copyright © 2020 Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
https://doi.org/10.11646/zootaxa.4868.1.4
http://zoobank.org/urn:lsid:zoobank.org:pub:DC4B8EC5-CBDA-49D2-B370-381C96AF3A10
Taxonomy of Peruvian water beetles in the genus Hydrochus Leach, 1817

(Coleoptera: Hydrochidae)
PHILIP D. PERKINS
Department of Entomology, Museum of Comparative Zoology, Harvard University, Cambridge, MA 02138 USA
� perkins@oeb.harvard.edu; https://orcid.org/0000-0001-8834-9052
Abstract
The taxonomy of Peruvian water beetles in the genus Hydrochus Leach, 1817 is presented. The known fauna now consists
of 22 species, 14 of which are described as new species. The new species are diagnosed, and high-resolution images of
the habitus and male genitalia of each species are given. The following new species are described: H. alternatus n. sp.,
H. astrictus n. sp., H. bisinuatus n. sp., H. bucapitus n. sp., H. cracentis n. sp., H. dualis n. sp., H. lescheni n. sp., H.
obliquus n. sp., H. parvicollis n. sp., H. spinosus n. sp., H. subovatus n. sp., H. tambopaticus n. sp., H. tripartitus n. sp.,
and H. uniformis n. sp. New locality data are provided for H. argutus Knisch 1921, H. bituberculatus Perkins 2019, H.
coeneni Makhan 1992, H. leei Perkins 2019, H. lobatus Perkins 2019, H. obscurus Sharp 1882, H. richteri Bruch 1915,
and H. sagittarius Perkins 2019. Images of the habitus and male genitalia of H. coeneni Makhan 1992 are also given.
Key words: Peru, water beetles, Hydrochidae, new species
Introduction
This is the sixth contribution in a series of papers revising the Neotropical species of the genus Hydrochus Leach,
1817. Previous papers have designated lectotypes (Perkins, 2019a, 2020a), described a remarkable new species
found in Brazil, Guyana and Suriname (Perkins, 2019b), and revised the Venezuelan, Guatemalan, and Ecuadorian
species (Perkins 2019c, 2020a, b, respectively). Revisions of Hydrochus from other Neotropical countries are in
progress.
Here, knowledge of the Hydrochus fauna of Peru is increased, based on the examination and study of more than
400 specimens, which are deposited in six institutions. The known fauna now consists of 22 species, 14 of which
are described as new species.
In addition to the description of new species, one of the goals of the revisions is to critically review, and correct
when necessary, prior taxonomic decisions and associated species distributions made in the works of D. Makhan
and A. Oliva. For background on the taxonomic and nomenclatural problems peculiar to Neotropical Hydrochus,
especially as they pertain to the publications of D. Makhan (1992, 1993, 1994, 1995a, b, 1996, 1998, 2000, 2001,
2004a, 2004b, 2005), readers are referred to Oliva (2000), Jäch (2006), Short & Hebauer (2006), Worthington et al.
(2016), and Perkins (2019a, c; 2020a, b).
Materials and methods
The materials and methods used for specimen preparation and specimen imaging follow those of Perkins (2019c).
Males cannot be differentiated from females on generally observable external characters. However, if the elytra are
raised, exposing the last genital tergite, females have a small to moderately large hairy lobe on each side, which is
absent in males (see figures of H. spanglerorum genital segments in Perkins (2019b).
Accepted by A. Short: 1 Oct. 2020; published: 23 Oct. 2020 61

Depositories
HNHM Hungarian Natural History Museum, Budapest, Hungary (G. Makranczy)

MCZ Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA (P. D. Perkins)
NMNH National Museum of Natural History, Smithsonian Institution, Washington, D.C., USA (C. J. Micheli)
NMW Naturhistorisches Museum, Wien, Austria (M. A. Jäch)
RBINS Institut royal des Sciences naturelles de Belgique, Brussels, Belgium (P. Limbourg)
SEMC Snow Entomological Museum, University of Kansas, Lawrence, Kansas, USA (A. E. Z. Short, Z. Falin)
Abbreviations for characters on male genitalia figures
The following abbreviations are used for male genitalia characters on the figures. More context concerning these
and other Hydrochus genitalia characters are present in Perkins (2019c).
Aedeagus characters:
adbl: aedeagus dorsal basal lobe

adm: aedeagus dorsal margin
adtl: aedeagus dorsal transition line (dorsal area where sclerotized and membranous parts of the aedeagus meet)
adm: aedeagus dorsal margin
agw: aedeagus greatest width
alm: aedeagus lateral margin
alw: aedeagus least width
aslw: aedeagus shaft least width (shaft = aedeagus part that is distal to adbl)
avm: aedeagus ventral margin
asgw: aedeagus shaft greatest width (shaft = aedeagus part that is distal to adbl)
avtl: aedeagus ventral transition line (ventral area where sclerotized and membranous parts
of the aedeagus meet)
gp: gonopore
Paramere characters:
pdmm: paramere dorsal medial margin

pdm: paramere dorsal margin
plsa: paramere lateral subapical angle
pmsm: paramere medial sclerotized margin
pseu: pseudoparamere
pvm: paramere ventral margin
pvmm: paramere ventral medial margin
pwsa: paramere weakly sclerotized area
Abbreviations used in species descriptions
E elytra
HW maximum width of head, at eyes
P pronotum
PA width of anterior margin of pronotum
PB width of posterior margin of pronotum
62 · Zootaxa 4868 (1) © 2020 Magnolia Press PERKINS

New Species Descriptions
Hydrochus alternatus, new species

Figs. 1, 3
FIGURE 1. Dorsal habitus of Hydrochus species.
Taxonomy of Peruvian Hydrochus Zootaxa 4868 (1) © 2020 Magnolia Press · 63

FIGURE 2. Dorsal habitus of Hydrochus species.
Type Material. Holotype (male): “PERU: Madre de Dios; Rio Tambopata Res; 30 air km SW Pto. Maldonado, 290
m, 16-20 xi 1979, J. B. Heppner, subtropical moist forest” (NMNH). Paratypes (6): PERU: Madre de Dios, Rio
Tambopata Res; 30 air km SW Pto. Maldonado, elev. 290 m (13°0’S, 69°33’W), 16–20.xi.1979, leg. J. B. Heppner
(2 NMNH); same locality, 21–25.xi.1979, leg. J. B. Heppner (2 NMNH); same locality, 26–30.xi.1979, leg. J. B.
Heppner (2 NMNH).
Differential Diagnosis. Among Peruvian species, recognized by the combination of small size (ca. 2.08 mm),
the narrow body form, the elytra with sutural interval and interstriae 3, 5, 7 and 9 costate, and the male genitalia (Fig.
3; described below).
Description. Habitus as illustrated (Fig. 3). Size: holotype (length/width, mm): body (length to elytral apices)
2.08/0.78; head width 0.54; pronotum l/w 0.47/0.50; PA 0.49; PB 0.43; elytra 1.42/0.78. Body size medium (ca. 2.08
mm). Dorsum black, with very slight if any iridescence, elytra without black spots. Legs dark brown. Punctation on
elytra ca. 1–2x that of pronotum. Elytra interstriae ca. 1x strial puncture diameter. Interstria 9th more convex than
others, overhanging 10th interstria. Usual area of callus on 5th not raised.

FIGURE 3. Hydrochus alternatus n. sp., habitus and male genitalia of holotype.
Pronotum slightly wider than long, widest slightly behind anterior margin, narrowed at base, sides weakly sinu-
ate, denticulate; anterior margin slightly bisinuate; depressions deep, densely punctate.
Elytra with sutural interval and interstriae 3, 5, 7 and 9 costate; spaces between strial punctures narrow walls to
ca. 0.5x their diameter; apices rather sharply conjointly rounded in dorsal view, in lateral view with punctate sub-
marginal groove, outer margin of apices without angulation.
Ventral characters: Mentum sparsely punctate, with median fovea. Submentum with two foveae.
Male genitalia (Fig. 3) general characters: genitalia elongate, about four times as long as widest part in dorsal/
ventral views; parameres in distal ½ much narrower than aedeagus, in proximal ½ ca. as wide as aedeagus, paramere

tips arcuate toward midline; pseudoparameres ca. ½ as long as parameres, wide proximally, narrowing distally and
with spinose margins apically and medially; aedeagus widest subapically, becoming narrower from agw to basal
area; basal piece shorter than parameres, distal ½ wider in dorsal/ventral views than lateral view, in dorsal/ventral
views markedly narrowed at proximal end, in lateral view arcuate and only slightly narrower at proximal end than
at distal end, orifice strongly sclerotized.
Dorsal surface: pdmm in proximal ½ slightly separated from alm, in distal ½ contiguous with, or very slightly
overlapping alm; adbl and adtl lacking.
Ventral surface: pvmm contiguous with, or only very narrowly separated from alm at about midlength, proximal
part narrowly separated from alm; pseudoparameres as described above.
Etymology. Named in reference to the costate odd-numbered elytral intervals.
Hydrochus astrictus, new species

Figs. 1, 4
Type Material. Holotype (male): “PERU: Tambopata Prov., Madre de Dios Dpto, 15km NE Puerto Maldonado,
Reserva Cuzco Amazonico, 12°33’S, 69°3’W, 200m, camp, 9 June 1989, J. S. Ashe, R. A. Leschen, #009, ex., at
light” (SEMC). Paratypes (35): PERU: Loreto, Pucallpa, 10–12.iv.1969, leg. P. & P. Spangler (11 NMNH); Madre
de Dios, Rio Tambopata Res; 30 air km SW Pto. Maldonado, elev. 290 m (13°0’S, 69°33’W), 2–5.xi.1979, leg. J.
B. Heppner (4 NMNH); same locality, 6–10.xi.1979, leg. J. B. Heppner (2 NMNH); same locality, 11–15.xi.1979,
leg. J. B. Heppner (4 NMNH); same locality, 16–20.xi.1979, leg. J. B. Heppner (8 NMNH); same locality, 21–
25.xi.1979, leg. J. B. Heppner (3 NMNH); same locality, 26–30.xi.1979, leg. J. B. Heppner (3 NMNH).
Differential Diagnosis. Among known Peruvian species, differentiated from all, except H. bisinuatus, by the
deep cervical groove behind the eyes. This species is a member of the H. collaris group (sensu Perkins 2019c); dif-
ferentiated from H. bisinuatus by the larger body size (ca. 3.00–3.38 vs. 2.74 mm), and the more strongly arcuate
anterior margin of the pronotum. However, reliable determinations will require careful examination of the male
genitalia (Fig. 4; described below).
3.16/1.10; head width 0.80; pronotum l/w 0.76/0.71; PA 0.71; PB 0.61; elytra 2.08/1.10. Dorsum grey to silver,
with moderately strong iridescence, cervical groove black, elytra with black spots. Legs testaceous to brown, with
tibiofemoral joints darker. Punctation on elytra ca. 2x that of pronotum. Elytra interstriae ca. 1–2x strial puncture
diameter. Interstria 9th more convex than others, overhanging 10th interstria. Usual area of callus on 5th moderately
raised, moderately elongate, ca. 6 punctures in row on each side.
Head with deep curving cervical groove from side to side behind eyes, groove punctate, cervical area behind
groove impunctate, shining, with very fine, low, closely set longitudinal raised lines.
Pronotum longer than wide (as ca. 76/71), widest at anterior margin, narrowed at base, sides straight or ex-
tremely slightly arcuate, smooth; anterior margin sinuate, median ½ markedly arcuate, fitting into cervical groove;
anterior angles very slightly produced; depressions moderately deep, more densely punctate than reliefs between
depressions.
Elytra with spaces between strial punctures ca. 0.5–1x their diameter; apices rather sharply conjointly rounded
in dorsal view, in lateral view outer margin with small angulation, impunctate area of apices rather large.
Ventral characters: Mentum coarsely densely punctate, with large, deep median fovea. Submentum with two
deep foveae, posterior margin arcuate.
Male genitalia (Fig. 4) general characters: shaft of genitalia long and moderately narrow, basal piece slightly
shorter than parameres; paramere tips slightly narrower than aedeagus shaft, medial margins strongly sclerotized,
remainder of tips very weakly sclerotized; aedeagus widest at adbl, shaft moderately narrow, margins sinuate, shaft
widest slightly before midlength (asgw), then narrowed at about proximal 1/3 of shaft (aslw), then becoming wider
before tapering at tip; gonopore on both dorsal and ventral surfaces, the latter much larger than the former; basal
piece almost cylindrical, with rim of orifice strongly sclerotized and distinctively hook-shaped.
Dorsal surface: pdmm closely approximating alm, except at adbl; adtl at about midlength of shaft; triangular
weakly sclerotized area of paramere very prominent; adbl very large.

FIGURE 4. Hydrochus astrictus n. sp., habitus and male genitalia of holotype.
Ventral surface: pvmm approximating alm except becoming very narrowly separated from one another and
overlapping shaft in proxima1 1/3; gonopore long, distinct, with sclerotized margins in proximal ½.
Etymology. Named in reference to the tightly fitting components of the head and prothorax: cervical groove
and anterior margin of the pronotum.

Hydrochus bisinuatus, new species
Figs. 1, 5
Type Material. Holotype (male): PERU: Loreto Prov., On the Amazon nr. the Shishista River, 400 m, 1 May 1992,
leg. James Danoff-Berg, ex: at lights (SEMC). Paratypes (4): PERU: Ucayali, Rio Calleria, 15 km from Ucalagi,
3–13.x.1961, leg. Borys Malkin (3 NMNH); Pucallpa, 10–12.iv.1969, leg. P. & P. Spangler (1 NMNH).
FIGURE 5. Hydrochus bisinuatus n. sp., habitus and male genitalia of holotype.

Differential Diagnosis. Among known Peruvian species, differentiated from all, except H. astrictus, by the
deep cervical groove behind the eyes. This species is a member of the H. collaris group (sensu Perkins 2019c);
differentiated from the known other species in the group by the more sinuate anterior margin of the pronotum, and
the smaller body size (ca. 2.74 mm). However, reliable determinations will require careful examination of the male
genitalia (Fig. 5; described below).
2.74/0.94; head width 0.68; pronotum l/w 0.32/0.33; PA 0.60; PB 0.49; elytra 1.66/0.94. Dorsum grey to silver, with
moderately strong iridescence, elytra with black spots. Legs testaceous to brown, with tibiofemoral joints darker.
Punctation on elytra ca. 2x that of pronotum. Elytra interstriae ca. 1–2x strial puncture diameter. Interstria 9th more
convex than others, overhanging 10th interstria. Usual area of callus on 5th well raised, moderately elongate, ca. 6
punctures in row on each side.
Head with deep curving cervical groove from side to side behind eyes, groove punctate, cervical area behind
groove impunctate, shining, with very fine, low, closely set longitudinal raised lines.
Pronotum very slightly wider than long (as ca. 30/31), widest at anterior margin, narrowed at base, sides slightly
sinuate, smooth; anterior margin sinuate, median ½ markedly arcuate, fitting into cervical groove; anterior angles
slightly produced; depressions deep to moderately deep, much more densely punctate than reliefs between depres-
sions, latter almost entirely impunctate.
Elytra with spaces between strial punctures ca. 1x their diameter; apices rather sharply conjointly rounded in
dorsal view, in lateral view outer margin with distinct angulation, impunctate area of apices rather large.
Ventral characters: Mentum coarsely densely punctate, with median fovea. Submentum with two deep foveae,
posterior margin arcuate.
Male genitalia (Fig. 5) general characters: genitalia elongate, about six times as long as widest part, basal piece
slightly shorter than parameres; parameres in distal ½ much narrower than aedeagus, in proximal ½ slightly wider
than aedeagus, paramere tips arcuate toward midline; aedeagus widest subapically, becoming narrower from agw to
basal area; basal piece slightly wider in dorsal/ventral views than lateral view, in lateral view very slightly arcuate,
lacking any process of sclerotized rim.
Dorsal surface: pdmm contiguous with, or only very narrowly separated from alm; adbl small; adtl very indis-
tinct, if present.
Ventral surface: pvmm contiguous with, or only very narrowly separated from alm in distal ¾, proximal part
overlapping alm slightly.
Etymology. Named in reference to the bisinuate anterior margin of the pronotum.
Hydrochus bucapitus, new species

Figs. 1, 6
Type Material. Holotype (male): “PERU: Madre de Dios; Rio Tambopata Res; 30 air km SW Pto. Maldonado,
290 m, 26–30 xi 1979, J. B. Heppner, subtropical moist forest” (NMNH). Paratype: Same data as holotype, except
5 xi 1979 (1 NMNH).
Differential Diagnosis. Differentiated from other Peruvian species by the combination of the moderate body
size (ca. 2.50 mm), the rather large head (width greater than width or length of pronotum), and the male genitalia
(Fig. 6; described below). Completely reliable determinations will include dissection of males.
2.50/0.83; head width 0.62; pronotum l/w 0.58/0.55; PA 0.55; PB 0.42; elytra 1.60/0.83. Body size moderate (ca.
2.50 mm), very narrow, except head relatively wide. Dorsum dark brown with slight iridescence, elytra without
black spots. Dorsum with numerous microsetae. Legs testaceous, with tibiofemoral joints and basal ½ of femora
darker. Punctation on elytra ca. 2–4x that of pronotum. Elytra interstriae narrow walls to ca. 0.5x strial puncture
diameter. Interstria 9th more convex than others, overhanging 10th interstria. Usual area of callus on 5th very weakly
raised.
Pronotum very slightly longer than wide, widest at anterior margin, narrowed at base, sides very weakly ar-
cuate, almost straight, smooth; anterior margin arcuate; depressions moderately deep, coarsely densely punctate,
reliefs between depressions less punctate.

Elytra with spaces between strial punctures narrow walls to ca. 0.5x their diameter; apices rather sharply con-
jointly rounded in dorsal view, in lateral view outer margin without angulation.
Ventral characters: Mentum moderately densely finely punctate, with large, deep median fovea. Submentum
with two large deep foveae.
Male genitalia (Fig. 6) general characters: genitalia very small, very slender, length ca. 6x width; parameres
pointed in dorsal/ventral views and lateral view, only slightly longer than aedeagus, lacking plsa; aedeagus narrowly
lanceolate, agw at about midlength; basal piece slightly shorter than parameres, width ca. equal in dorsal/ventral and
lateral views, orifice sclerotized, inverted V-shape in dorsal/ventral views.
FIGURE 6. Hydrochus bucapitus n. sp., habitus and male genitalia of holotype.

Dorsal surface: adbl large; pdmm approximating alm.
Ventral surface: pvmm over middle ½ overlapping aedeagus, margins very narrowly separated one from the
other.
Etymology. Named in reference to the relatively large head.
Hydrochus cracentis, new species

Figs. 1, 7
FIGURE 7. Hydrochus cracentis n. sp., habitus and male genitalia of holotype.

light” (SEMC).
Differential Diagnosis. Differentiated from other Peruvian species by the combination of the small body size
(ca. 2.11 mm), the narrow body form (width/length ca. 0.33), the numerous dorsal microsetae, and the male genitalia
(Fig. 7; described below). Completely reliable determinations will include dissection of males.
2.11/0.71; head width 0.54; pronotum l/w 0.49/0.47; PA 0.47; PB 0.40; elytra 1.33/0.71. Body size small (ca. 2.11
mm), very narrow. Dorsum brown to copper (holotype teneral), with slight to moderately strong iridescence, elytra
with black spots. Dorsum with numerous microsetae. Legs testaceous, with tibiofemoral joints and basal ½ of femora
darker. Punctation on elytra ca. 2–3x that of pronotum. Elytra interstriae ca. 0.5–1x strial puncture diameter. Interstria
9th more convex than others, overhanging 10th interstria. Usual area of callus on 5th very weakly if at all raised.
Pronotum very slightly longer than wide, widest at anterior margin, narrowed at base, sides very weakly ar-
cuate, almost straight, smooth; anterior margin arcuate; depressions moderately deep, coarsely densely punctate,
reliefs between depressions less punctate.
in dorsal view, in lateral view outer margin without angulation.
Ventral characters: Mentum densely punctate, with median fovea. Submentum with two foveae.
Male genitalia (Fig. 7) general characters: genitalia very small, wider in dorsal/ventral views than lateral view;
paramere tips extending slightly beyond tip of aedeagus, in dorsal/ventral views slightly widened, arcuate toward
midline, with apical portions separated one from the other, in lateral view tips straight, distal end rounded; aedeagus
broad in both dorsal/ventral and lateral views, in distal ½ much wider than parameres in dorsal/ventral views; basal
piece longer and slightly wider than parameres; basal piece much wider in dorsal/ventral views than lateral view.
Dorsal surface: pdmm slightly separated from alm in distal portion, contiguous or nearly so for remainder of length.
Ventral surface: ventrally parameres in distal ½ distinctly wider than dorsal part of parameres, pvmm in this area
very thin and nearly straight, overlapping alm; pvmm in proximal ½ becoming narrowly separate one from the other
and with thin, finely striated ridge; pseu small, appear to be between parameres and aedeagus.
Etymology. Named in reference to the slender body form.
Hydrochus dualis, new species

Figs. 1, 8
Type Material. Holotype (male): “Peru, S. A. Dec. 11–30, 1937, F. Woytkowski No. 3811 // Department Huanuco,
Vic. Leonpampa, jungle, 800 m. a. s. l.” (SEMC).
Differential Diagnosis. Among Peruvian species, recognized by the combination of large size (ca. 3.92 mm),
the moderately deep pronotal depressions, the roundly raised, almost impunctate pronotal reliefs, the 5th elytral in-
terstria with two callosities, the usual posterior and another, smaller, slightly anterior, and the male genitalia (Fig. 8;
described below). Completely reliable determinations will include dissection of males.
3.92/1.54; head width 0.90; pronotum l/w 0.87/0.92; PA 0.82; PB 0.77; elytra 2.54/1.54. Body size large (ca. 3.92
mm). Dorsum grey with some black on head, with slight to moderate iridescence, elytra with black spots. Legs
brown, with tibiofemoral joints darker. Punctation on elytra ca. 0.5–1x that of pronotum. Elytra interstriae ca. 1–3x
strial puncture diameter. Interstria 9th more convex than others, overhanging 10th interstria. Usual area of callus on
5th markedly raised, moderately elongate, ca. 5 punctures in row on each side.
Pronotum very slightly wider than long, widest slightly behind anterior margin, narrowed at base, sides arcu-
ate or very weakly sinuate, smooth; anterior margin slightly bisinuate; depressions moderately deep, more dense-
ly punctate than low reliefs between depressions, latter almost entirely impunctate; anteromedian depression less
punctate than others.
Elytra with spaces between strial punctures ca. 1–2x their diameter; apices rather sharply conjointly rounded in
dorsal view, in lateral view outer margin with moderate angulation; 3rd and 4th interstriae slightly raised immediately
anterior to usual 5th callosity; 5th interstria with two callosities, the usual posterior and another, smaller, slightly
anterior.

Ventral characters: Mentum densely punctate, with large, deep median fovea. Submentum with two foveae.
Male genitalia (Fig. 8) general characters: genitalia very stout, wide in lateral view; paramere tips extending
slightly beyond distal end of aedeagus, lateral margin between plsa and distal end emarginate, medial margin arcu-
ate, in lateral view extreme end of tips upturned slightly; aedeagus lanceolate, greatest width distal to midlength,
with alsm; pseudoparameres large, extending to level of distal end of aedeagus, pseudoparameres wider in lateral
than dorsal/ventral views; basal piece shorter than parameres, orifice damaged during dissection.
Dorsal surface: adtl located proximal to midlength of aedeagus; pdmm sinuate, overlapping lateral margins of
aedeagus only over proximal 1/4; adbl moderately large.
Ventral surface: avtl U-shaped, near basal 1/5 of aedeagus; pvmm not overlapping lateral margins of aedeagus;
pseudoparameres enveloping most of basal ¼ of parameres.
FIGURE 8. Hydrochus dualis n. sp., habitus and male genitalia of holotype.
Etymology. Named in reference to the two callosities on the 5th interstria.

Remarks. The male genitalia of H. dualis are similar in some characters to that of H. ramcharani Makhan (fig-
ure of aedeagus given in Perkins, 2019c). Differences in the two species include: 1) in H. dualis the paramere tips,

from plsa to distal end, are shorter and the lateral margin is entirely emarginate, whereas in H. ramcharani the tips
are nearly twice as long and the lateral margin is straight except emarginate subapically; 2) in H. dualis the pseudo-
parameres are much wider in lateral view for the entire length, whereas in H. ramcharani the pseudoparameres are
cylindrical beyond ca. the basal 1/3; and 3) in H. dualis the adtl is located more proximally than in H. ramcharani,
hence the membranous dorsal part of the aedeagus is much larger in H. dualis.
Hydrochus lescheni, new species

Figs. 2, 9
light” (SEMC).
the shallow pronotal depressions, more densely punctate than the low reliefs between depressions, latter almost en-
tirely impunctate, and the male genitalia (Fig. 9; described below). Completely reliable determinations will include
dissection of males.
2.12/0.82; head width 0.59; pronotum l/w 0.52/0.55; PA 0.50; PB 0.44; elytra 1.29/0.82. Dorsum grey to silver grey,
with slight to moderately strong iridescence, elytra with black spots (holotype specimen teneral). Legs testaceous,
with tibiofemoral joints and basal ½ of femora darker. Punctation on elytra ca. 2–3x that of pronotum. Elytra inter-
striae ca. 1–1.5x strial puncture diameter. Interstria 9th more convex than others, overhanging 10th interstria. Usual
area of callus on 5th very weakly raised, moderately elongate, ca. 6 punctures in row on each side.
Pronotum length slightly less than width, widest slightly behind anterior margin, narrowed at base, sides arcuate
or very weakly sinuate, smooth; anterior margin slightly bisinuate; depressions shallow, more densely punctate than
low reliefs between depressions, latter almost entirely impunctate.
in dorsal view, in lateral view outer margin without angulation.
Ventral characters: Mentum densely punctate, with large median fovea. Submentum with two large foveae.
Male genitalia (Fig. 9) general characters: genitalia very small; paramere tips extending well beyond tip of ae-
deagus, in dorsal/ventral views slightly widened, with apical portions well separated one from the other, and medial
margins nearly parallel, in lateral view tips narrowing apically, slightly arcuate toward ventral, extreme tip with
pointed apex; aedeagus broad in both dorsal/ventral and lateral views, much wider than parameres in dorsal/ventral
views, internally in central portion with many concentric lines; basal piece shorter than parameres, widened at ori-
fice in dorsal/ventral views, in lateral view basal piece markedly narrowed from distal end to base.
Dorsal surface: pdmm slightly overlapping alm.
Ventral surface: pvmm slightly overlapping alm for most of length, becoming more narrowly separated from
one another basally.
Etymology. Named in honor of the collector, Richard A. Leschen.
Hydrochus obliquus, new species

Figs. 1, 10
Type Material. Holotype (male): “PERU: Dept. Huanuco, Rio Llullapichis, Panguana, 260 m a.s.l. 9°37’S,
74°56’W, 18.11–16.12.09, leg. Wachtel” (NMW).
the black dorsum, the densely, deeply punctate pronotum with deep depressions, the elytra with sutural and intervals
3, 5, 7, and 9 costate, 5th and 9th more strongly raised than others, the spaces between strial punctures narrow walls
to ca. 0.5x their diameter, producing zigzag pattern of intervals, and the male genitalia (Fig. 10; described below).
mm). Dorsum black, with slight to moderately strong iridescence, elytra without black spots. Legs brown, lateral

margin of tibiae with row of spines. Head with frons medially slightly tumid, with longitudinal punctate groove
on each side of tumidity. Punctation on elytra ca. 2–5x that of pronotum. Elytra interstriae narrow walls to ca. 0.5x
5th not raised.
Pronotum densely deeply punctate, slightly longer than wide, widest at anterior margin, narrowed at base, sides
arcuate or very weakly sinuate, smooth; anterior margin slightly bisinuate; depressions deep, slightly more densely
punctate than reliefs between depressions.
FIGURE 9. Hydrochus lescheni n. sp., habitus and male genitalia of holotype.

Elytra with sutural and intervals 3, 5, 7, and 9 costate, 5th and 9th more strongly raised than others; spaces be-
tween strial punctures narrow walls to ca. 0.5x their diameter, producing zigzag pattern of intervals; apices rather
sharply conjointly rounded in dorsal view, in lateral view outer margin without angulation.
Ventral characters: Mentum moderately coarsely and moderately densely punctate, with small shallow median
fovea. Submentum narrow, without foveae. Legs brown, lateral margin of each tibia with row of prominent spines.
Male genitalia (Fig. 10) general characters: genitalia without parameres; basal piece elongate, arcuate in lateral view,
extreme distal end bifurcate; distal piece emerging from bifurcation, long, straight, tapering apically, lacking processes.
FIGURE 10. Hydrochus obliquus n. sp., habitus and male genitalia of holotype.

Etymology. Named in reference to the zigzag pattern of the elytral intervals.
Remarks. Makhan (1998, 2001) proposed several new genera in the Hydrochidae. Hansen (1999), Short & He-
bauer (2006), and Short & Fikáček (2011) have considered these to be invalid, and have placed them in synonymy
with Hydrochus. Based on the male genitalia, H. obliquus n. sp. is related to species placed in one of Makhan’s
“genera.” The taxonomy of the very diverse genus Hydrochus would benefit from a comprehensive molecular
phylogenetic study. Until such time, the following statement from Hansen (1999) seems to hold true: “Hence, there
seems to be no justification for considering [Makhan’s “genera”] as distinct genera … there is certainly no phyloge-
netic support for generic rank (this would almost inevitably leave Hydrochus as paraphyletic).”
Hydrochus parvicollis, new species

Figs. 1, 11
Type Material. Holotype (male): “PERU: Madre de Dios; Rio Tambopata Res; 30 air km SW Pto. Maldonado,
290 m, 6–10 xi 1979, J. B. Heppner, subtropical moist forest” (NMNH). Paratypes (4): Same data as holotype (4
NMNH).
the rather small pronotum, head width greater than pronotum width or length, and the male genitalia (Fig. 11; de-
scribed below). Completely reliable determinations will include dissection of males.
2.31 mm). Dorsum grey to bright silver metallic, with moderately strong iridescence, elytra with several black spots.
Legs brown to testaceous, with tibiofemoral joints darker. Punctation on elytra deep, ca. 2–3x that of pronotum.
Elytra interstriae ca. 1x strial puncture diameter. Interstriae 5th and 9th more convex than others, 9th overhanging 10th
interstria. Usual area of callus on 5th very slightly raised, if at all.
Pronotum slightly wider than long, widest slightly behind anterior margin, markedly narrowed at base, sides
very slightly sinuate, smooth; anterior margin bisinuate; depressions moderately deep, much more densely punctate
than reliefs.
Elytra with spaces between strial punctures narrow ridges to ca. 0.5x their diameter; apices rather sharply sepa-
rately rounded in dorsal view, in lateral view outer margin with very slight angulation.
Ventral characters: Mentum densely punctate, with median fovea. Submentum without foveae.
Male genitalia (Fig. 11) general characters: Male genitalia very small; parameres apically arcuate in ca. 90
degree angle medially directed, distal tips pointed, with pmsm in distal 1/2; aedeagus in dorsal/ventral views with
distal 2/3 much wider than parameres, distal end wide and apically truncate, agw at ca. distal 1/3, width narrowed
over proximal 1/4; basal piece shorter than parameres, in dorsal/ventral views tapering from distal to base, arcuate
in lateral view, orifice damaged during dissection.
Dorsal surface: pdmm overlapping aedeagus basally.
Ventral surface: pvmm produced in lightly sclerotized lobe with minute ridges, overlapping aedeagus
Etymology. Named in reference to the small prothorax, in relation to the size of the head.
Hydrochus spinosus, new species

Figs. 2, 12
light” (SEMC).
the shallow pronotal depressions, more densely punctate than the low reliefs between depressions, latter almost
entirely impunctate, the elytral punctures much larger than the pronotal punctures, and the male genitalia (Fig. 12;
described below). Completely reliable determinations will include dissection of males.

FIGURE 11. Hydrochus parvicollis n. sp., habitus and male genitalia of holotype.
mm), narrow. Dorsum light brown with slight iridescence (holotype is teneral), elytra without black spots. Dorsum
with numerous microsetae. Legs testaceous, with tibiofemoral joints darker. Punctation on elytra ca. 3–5x that of
pronotum. Elytra interstriae narrow walls to ca. 0.5x strial puncture diameter. Interstria 9th more convex than others,
overhanging 10th interstria. Usual area of callus on 5th very weakly raised.
Pronotum length and width equal, widest at anterior margin, narrowed at base, sides very weakly arcuate,
almost straight, smooth; anterior margin arcuate; depressions shallow, coarsely densely punctate, reliefs between
depressions almost impunctate.
Elytra with spaces between strial punctures narrow walls to ca. 0.5x their diameter; apices rather sharply con-
jointly rounded in dorsal view, in lateral view outer margin without angulation.

FIGURE 12. Hydrochus spinosus n. sp., habitus and male genitalia of holotype.
Ventral characters: Mentum moderately densely punctate, with small, deep median fovea. Submentum large,
with two large deep foveae.
Male genitalia (Fig. 12) general characters: genitalia very small; parameres much longer than aedeagus, tips
arcuate toward midline, distal ends pointed; four pseudoparameres, two short and two much longer, the latter with
numerous short spines on distal ends; aedeagus widest subapically, basal 2/3 parallel sided or nearly so; basal piece
much shorter than parameres, oval in dorsal/ventral views, much narrower in lateral view.
Dorsal surface: aedeagus overlapping pdmm.
Ventral surface: medial margins of large pseudoparameres overlapping lateral margin of aedeagus.
Etymology. Named in reference to the spinose pseudoparameres.

Hydrochus subovatus, new species
Figs. 1, 13
FIGURE 13. Hydrochus subovatus n. sp., habitus and male genitalia of holotype.
Type Material. Holotype (male): “PERU: Madre de Dios, Tambopata, Terrafirme, Drinking Quebrada, (13°0’S,
69°33’W) // 11.ix.1983, leg. J. J. Anderson, ex leaf packs” (NMNH). Paratype: Same data as holotype (1 NMNH).
the sub-oval body form, the elytra with odd-numbered interstriae raised, and the male genitalia (Fig. 13; described
below).

2.30 mm). Dorsum dark brown, with slight iridescence, elytra without black spots. Legs brown, with tibiofemoral
joints and basal ½ of femora darker. Punctation on elytra ca. 2–3x that of pronotum. Elytra interstriae ca. 0.5–1x
5th very weakly raised, moderately elongate.
Pronotum slightly wider than long, widest slightly behind anterior margin, narrowed at base, sides arcuate or
very weakly sinuate, denticulate, each process with moderately long arcuate seta; anterior margin bisinuate; depres-
sions, especially anterior, deep, much more densely punctate than reliefs between depressions.
Elytra with odd numbered interstriae raised, spaces between strial punctures ca. 0.5–1x their diameter; apices
rather sharply conjointly rounded in dorsal view, with sublateral punctate groove, in lateral view outer margin with
slight angulation.
Ventral characters: Mentum moderately densely punctate, with median fovea. Submentum with two deep fo-
veae.
Male genitalia (Fig. 13) general characters: genitalia very small, very slender, length ca. 6x width; paramere dis-
tal ends sharply rounded in dorsal/ventral and lateral views, slightly longer than aedeagus, lacking plsa, tips beyond
aedeagus with medial margins parallel or nearly so, distal ½ of parameres with distinct pmsm; aedeagus narrowly
lanceolate, agw slightly distal to midlength, alw just distal to adbl; basal piece slightly longer than parameres, width
slightly greater in dorsal/ventral than lateral views, orifice weakly sclerotized.
Dorsal surface: adbl moderately large, almost as wide as agw; pdmm approximating shape of alm.
Ventral surface: pvmm basally overlapping aedeagus, margins very narrowly separated one from the other,
otherwise approximating shape of alm.
Etymology. Named in reference to the oval habitus.
Hydrochus tambopaticus, new species

Figs. 1, 14
m, 2–5 xi 1979, J. B. Heppner, subtropical moist forest” (NMNH). Paratypes (75): PERU: Same data as holotype (4
NMNH); same locality, 6–10.xi.1979, leg. J. B. Heppner (9 NMNH); same locality, 11–15.xi.1979, leg. J. B. Heppner
(15 NMNH); same locality, 16–20.xi.1979, leg. J. B. Heppner (5 NMNH); same locality, 21–25.xi.1979, leg. J. B.
Heppner (4 NMNH); Ucayali, Rio Calleria, 15 km from Ucalagi, 3–13.x.1961, leg. Borys Malkin (1 NMNH); Loreto,
Pucallpa, 10–12.iv.1969, leg. P. & P. Spangler (8 NMNH); Madre de Dios, Parque Manu, Pakitza, Trocha Dos, c.53,
berlesed leaf litter, elev. 250 m (12°7’S, 70°58’W), 13.ix.1989, leg. R. A. Faitoute (39) (3 NMNH); same locality,
19.ix.1989, leg. R. A. Faitoute (47a) (2 NMNH); same locality, 20.ix.1989, leg. R. A. Faitoute (48) (4 NMNH).
Differential Diagnosis. Among Peruvian species, recognized by the combination of large size (ca. 3.85–3.93
mm), the shallow pronotal depressions, the large elytral tubercles, and the male genitalia (Fig. 14; described below).
This species has a generalized Hydrochus habitus; reliable determinations will require dissection of males. The male
genitalia are similar in general characters to that of H. elsjeae Makhan from Bolivia, but differ in several characters
including 1) larger size (length as ca. 5.60 to 4.35); 2) aedeagal apex more nearly attaining apex of parameres; 3)
paramere tips with psla more acute and lateral margins less strongly emarginate; 4) pseudoparameres much wider in
ventral view; 5) genitalia wider in lateral view.
3.93/1.43; head width 0.95; pronotum l/w 0.86/0.90; PA 0.83; PB 0.76; elytra 2.52/1.43. Body size large (ca. 3.93
mm). Dorsum grey to black, with slight iridescence, elytra with black spots. Legs testaceous to brown, with tibio-
femoral joints darker. Punctation on elytra ca. 0.5–1x that of pronotum. Elytra interstriae ca. 1–2x strial puncture
diameter. Interstria 9th more convex than others, overhanging 10th interstria. Usual area of callus on 5th markedly
raised, moderately elongate, ca. 5 punctures in row on each side.
Pronotum very slightly wider than long, widest slightly behind anterior margin, narrowed at base, sides arcu-
ate or very weakly sinuate, smooth; anterior margin arcuate; depressions shallow, more densely punctate than low
reliefs between depressions, latter almost entirely impunctate; anteromedian depression less punctate than others.

FIGURE 14. Hydrochus tambopaticus n. sp., habitus and male genitalia of holotype.
Elytra with spaces between strial punctures ca. 1x their diameter; apices rather sharply conjointly rounded in
dorsal view, in lateral view outer margin with moderate angulation.
Ventral characters: Mentum densely punctate, with large, deep median fovea. Submentum with two foveae.
Male genitalia (Fig. 14) general characters: genitalia very stout, wide in lateral view; paramere tips extending
slightly beyond distal end of aedeagus, plsa acutely pointed, lateral margin between plsa and distal end slightly
emarginate, medial margin arcuate, in lateral view extreme end of tips upturned slightly; aedeagus lanceolate, great-
est width distal to midlength, with alsm; pseudoparameres large, not extending to level of distal end of aedeagus
or to plsa, pseudoparameres not wider in lateral than dorsal/ventral views; basal piece as long as parameres, and as
long as aedeagus (including adbl), orifice in dorsal/ventral views much narrower than greatest width of basal piece,
strongly sclerotized and, in lateral view, produced in rounded process.
Dorsal surface: adtl located slightly distal to midlength of aedeagus; pdmm sinuate, overlapping lateral margins
of aedeagus; adbl moderately large; gp large.

Ventral surface: avtl U-shaped, near midlength of aedeagus; pvmm overlapping lateral margin of aedeagus only
at midlength; pseudoparameres wide basally, narrowed apically and with minute spines.
Etymology. Named in reference to the geographical distribution.
Hydrochus tripartitus, new species

Figs. 1, 15
m, 26–30 xi 1979, J. B. Heppner, subtropical moist forest” (NMNH). Paratypes (17): Peru: Same data as holotype
(3 NMNH); Madre de Dios, Rio Tambopata Res; 30 air km SW Pto. Maldonado, elev. 290 m (13°0’S, 69°33’W),
6–10.xi.1979, leg. J. B. Heppner (1 NMNH); same locality, 11–15.xi.1979, leg. J. B. Heppner (2 NMNH); same lo-
cality, 16–20.xi.1979, leg. J. B. Heppner (7 NMNH); same locality, 21–25.xi.1979, leg. J. B. Heppner (4 NMNH).
Differential Diagnosis. Among Peruvian species, recognized by the combination of small size (ca. 1.76–2.50
mm), the brown to black dorsum, the sub-oval body shape, the densely punctate pronotum, with deep depressions,
the elytra with sutural and intervals 3, 5, 7, and 9 costate, and the male genitalia (Fig. 15; described below).
Description. Habitus as illustrated (Fig. 15). Size: holotype (length/width, mm): body (length to elytral api-
ces) 1.76/0.72; head width 0.47; pronotum l/w 0.39/0.45; PA 0.43; PB 0.41; elytra 1.11/0.72. Body size small (ca.
1.76–2.30 mm). Dorsum dark brown to black, with slight to moderately strong iridescence, elytra without black
spots. Legs brown, lateral margin of mesotibiae with row of spines. Head with frons medially slightly tumid, with
longitudinal punctate groove on each side of tumidity. Punctation on elytra ca. 2–3x that of pronotum. Elytra inter-
striae ca. 0.5–1x strial puncture diameter. Interstria 9th more convex than others, overhanging 10th interstria. Usual
area of callus on 5th not raised.
Pronotum densely punctate, slightly wider than long, widest slightly behind anterior margin, narrowed at base,
sides weakly sinuate, smooth; anterior margin slightly bisinuate; depressions deep, slightly more densely punctate
than reliefs between depressions.
Elytra with sutural and intervals 3, 5, 7, and 9 costate; spaces between strial punctures narrow walls to ca. 0.5x their
diameter; apices rather sharply conjointly rounded in dorsal view, in lateral view outer margin without angulation.
Ventral characters: Mentum moderately coarsely and moderately densely punctate, with shallow median fovea.
Submentum narrow, without foveae.
Male genitalia (Fig. 15) general characters: genitalia without parameres, basal piece elongate, arcuate in lateral
view; distal piece arcuate in lateral view, terminating in three processes: short slender gonopore bearing flagellum,
small hook-shaped process, and larger blade-like process.
Etymology. Named in reference to the three processes of the distal piece of the male genitalia.
Remarks. Refer to the remarks section of H. obliquus, which also apply to H. tripartitus n. sp.
Hydrochus uniformis, new species

Figs. 1, 16
light” (SEMC).
the dense and deep dorsal punctation, the interstices with sparse to moderately dense microsetae, the pronotal punc-
tures of very uniform size, , and the male genitalia (Fig. 16; described below). Completely reliable determinations
will include dissection of males.
2.27 mm). Dorsum dark brown to black, with slight iridescence, elytra without distinct black spots. Legs testaceous,
with tibiofemoral joints and basal ½ of femora darker. Dorsal punctuation dense and deep. Punctation on elytra ca.
4–5x that of pronotum. Elytra interstriae narrow ridges forming zig-zag pattern to ca. 1x strial puncture diameter,

with sparse microsetae. Interstria 9th more convex than others, overhanging 10th interstria. Usual area of callus on 5th
weakly raised, moderately elongate, ca. 5 punctures in row on each side.
FIGURE 15. Hydrochus tripartitus n. sp., habitus and male genitalia of holotype.
Pronotum densely coarsely punctate, of quite uniform size; interstices rounded, with rather dense microsetae,
length less than width, widest slightly behind anterior margin, narrowed at base, sides arcuate or very weakly sinu-
ate, smooth; anterior margin slightly bisinuate; anterior depressions moderately deep.
Elytra with sutural interval raised over posterior declivity; spaces between strial punctures narrow walls to ca.
0.5x their diameter; apices slightly produced, separately rounded in dorsal view, in lateral view outer margin without
angulation.

FIGURE 16. Hydrochus uniformis n. sp., habitus and male genitalia of holotype.
Ventral characters: Mentum densely punctate, with large median fovea. Submentum with two large foveae.
Male genitalia (Fig. 16) general characters: short and rather broad, basal piece longer than parameres; paramere
tips extending to level of aedeagus tip, or very slightly beyond, weakly sclerotized, widened slightly, arcuate slightly
toward midline but not contacting one another; aedeagus wider than parameres, broad even at base; basal orifice
distinctively shaped, broad in lateral view, with apicoventral arcuate process.
Dorsal surface: pdmm contiguous with alm in distal ½, overlapping alm slightly in proximal ½; adtl and adbl absent.
Ventral surface: pvmm slightly overlapping alm; parameres fused to one another basally; pseudoparameres
large, ca. ½ length of paramere.
Etymology. Named in reference to the uniform size of the punctures on the head and pronotum.

New Distribution Records
Hydrochus argutus Knisch

Fig. 2
Hydrochus argutus Knisch 1921: 3.

Hydrochus johannapietersenae Makhan 1995; Oliva (1996); Perkins (2019a).
Originally described from Brazil. Also reported from Argentina, Bolivia, and Paraguay (Oliva, 1996); however, ad-
ditional confirmation of the records from Argentina and Bolivia are needed. Confirmation of the Paraguay record
(H. johannapietersenae jr. syn.) was given in Perkins (2019a). Locality and deposition data records have been pub-
lished for Ecuador (Perkins 2020b). Records from Peru are given below. Habitus and male genitalia illustrations are
given in Perkins (2019a).
Material examined (79): Peru: Tambopata, lodge, uv light, 14.viii.1982, leg. J. J. Anderson (1 NMNH); Ucay-
ali, Rio Calleria, 15 km from Ucalagi, 3–13.x.1961, leg. Borys Malkin (1 NMNH); Madre de Dios, Rio Tambopata
Res; 30 air km SW Pto. Maldonado, elev. 290 m (13°0’S, 69°33’W), 2–5.xi.1979, leg. J. B. Heppner (14 NMNH);
same locality, 6–10.xi.1979, leg. J. B. Heppner (16 NMNH); same locality, 11–15.xi.1979, leg. J. B. Heppner (8
NMNH); same locality, 16–20.xi.1979, leg. J. B. Heppner (9 NMNH); same locality, 21–25.xi.1979, leg. J. B. Hep-
pner (17 NMNH); same locality, 26–30.xi.1979, leg. J. B. Heppner (13 NMNH).
Hydrochus bituberculatus Perkins

Fig. 2
Hydrochus bituberculatus Perkins, 2019c: 11.
Originally described from Venezuela; locality data records also published for Ecuador (Perkins 2020b); records
from Peru are given below. Habitus and male genitalia illustrations are given in Perkins (2019c).
Material examined (96): PERU: Huanuco, Vic. Leonpampa, jungle, elev. 800 m, 11–30.xii.1937, leg. F.
Woytkowski (3811) (13 SEMC); Loreto, Pucallpa, 10–12.iv.1969, leg. P. & P. Spangler (22 NMNH); San Antonio,
1.viii.1965, leg. J. C. Hitchcock (2 NMNH); Madre de Dios, Rio Tambopata Res; 30 air km SW Pto. Maldonado,
elev. 290 m (13°0’S, 69°33’W), 2–5.xi.1979, leg. J. B. Heppner (27 NMNH); same locality, 6–10.xi.1979, leg. J. B.
Heppner (10 NMNH); same locality, 11–15.xi.1979, leg. J. B. Heppner (11 NMNH); same locality, 16–20.xi.1979,
leg. J. B. Heppner (2 NMNH); same locality, 21–25.xi.1979, leg. J. B. Heppner (5 NMNH); same locality, 26–
30.xi.1979, leg. J. B. Heppner (6 NMNH).
Hydrochus coeneni Makhan

Fig. 17
Hydrochus coeneni Makhan 1992: 102

Originally described from Brazil; new records from Peru are given below.
Material examined (8): Peru: Madre de Dios, Rio Tambopata Res; 30 air km SW Pto. Maldonado, elev. 290 m
(13°0’S, 69°33’W), 2–5.xi.1979, leg. J. B. Heppner (1 NMNH); same locality, 6–10.xi.1979, leg. J. B. Heppner (3
NMNH); same locality, 16–20.xi.1979, leg. J. B. Heppner (1 NMNH); same locality, 26–30.xi.1979, leg. J. B. Hep-
pner (1 NMNH); Tambopata, Madre de Dios, 15km NE Puerto Maldonado, Reserva Cuzco Amazonico, elev. 200 m
(12°33’S, 69°3’W), 9.vi.1989, leg. J. S. Ashe, R. A. Leschen (009) (1 SEMC); same locality, 14.vi.1989, leg. J. S.
Ashe, R. A. Leschen (180) (1 SEMC).
Remarks. I have examined the holotype, including the male genitalia, of H. coeneni (RBINS). The male geni-
talia has the tips of the parameres sheared off—by a previous worker. The remainder of the male genitalia, including
the basal piece, are distinctively shaped, allowing the identification. The male genitalia drawing of Makhan (1992)
shows the paramere tips to be present, but the drawing is very generalized, and only one view is given.

FIGURE 17. Hydrochus coeneni Makhan, habitus and male genitalia of non-type.
Hydrochus leei Perkins

Fig. 2
Hydrochus leei Perkins, 2019c: 50.
Originally described from Venezuela; locality data records also published for Guatemala (Perkins 2020a). The one
known record from Peru is given below. Habitus and male genitalia illustrations are given in Perkins (2019c).

Material examined (1): PERU: Loreto, Rio Yarapa, Puerto Miguel, elev. 200 m, 16–23.xii.1994, leg. T. Hatz
& G. Holzinger (1 HNHM).
Hydrochus lobatus Perkins

Fig. 2
Hydrochus lobatus Perkins, 2019c: 42.
Originally described from Venezuela; records from Peru are given below. Habitus and male genitalia illustrations
are given in Perkins (2019c).
Material examined (5): PERU: Madre de Dios, Rio Tambopata Res; 30 air km SW Pto. Maldonado, elev. 290
m (13°0’S, 69°33’W), 11–15.xi.1979, leg. J. B. Heppner (1 NMNH); same locality, 16–20.xi.1979, leg. J. B. Hep-
pner (1 NMNH); same locality, 21–25.xi.1979, leg. J. B. Heppner (1 NMNH); same locality, 26–30.xi.1979, leg. J.
B. Heppner (2 NMNH).
Hydrochus obscurus Sharp

Fig. 2
Hydrochus obscurus Sharp, 1882: 90
Originally described from Guatemala. Records from several South American countries are in the literature; how-
ever, some of these records are invalid synonymies proposed by Oliva (1992), for H. corruscans Bruch (Argentina)
(see Perkins 2019a), and H. elsjeae Makhan (Bolivia) (Perkins, in prep.). Locality data and deposition records have
been published for Venezuela (Perkins 2019c). The one locality data record from Peru is given below. Habitus and
male genitalia illustrations are given in Perkins (2019c, 2020a).
Material examined (3): PERU: Madre de Dios, Rio Tambopata Res; 30 air km SW Pto. Maldonado, elev. 290
m (13°0’S, 69°33’W), 16–20.xi.1979, leg. J. B. Heppner (3 NMNH).
Hydrochus richteri Bruch
Hydrochus richteri Bruch, 1915: 463; Oliva (1992) lectotype designation

Hydrochus soesilaae Makhan, 1994; syn. Oliva (1996)
Originally described from Argentina. This species is also recorded from Paraguay (Oliva 1996); however, this needs
confirmation as there is a synonymy dispute between Oliva (1996) and Makhan (1998). Two records from Peru are
given below.
Material examined (32): PERU: Loreto, 15km from Ucayali on R. Calleria, Colonia Calleria, 10–20.xi.1961,
leg. B. Malkin (1 NMNH); Pucallpa, Lago Yarina Cocha, 11.iv.1969, leg. P. & P. Spangler (31 NMNH).
Hydrochus sagittarius Perkins

Fig. 2
Hydrochus sagittarius Perkins, 2019c: 20
Originally described from Venezuela; the record from Peru is given below. Habitus and male genitalia illustrations
are given in Perkins (2019c).
Material examined (3): PERU: Cajamarca, Nanchoc Quebrada, elev. 400 m (6°57’S, 79°15’W), 21.xii.1984–
17.i.1985, leg. James M. Carpenter (3 MCZ).

Acknowledgments
The curators listed in the depositories section are sincerely thanked for kindly providing specimens of Hydrochus.
Special thanks to the following curators for providing loans of crucial type specimens, without which this study
would not have been possible: Max Barclay (NHM, London), Manfred Jäch (NMW, Vienna), Bernd Jaeger (ZMHB,
Berlin), Pol Limbourg (RBINS, Brussels), and György Makranczy (HNHM, Budapest).
References
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Perkins 2020 Escarabajos Buceadores de Tambopata 10.11646@zootaxa.4868.1.4

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Zootaxa 4868 (1): 061–089 ISSN 1175-5326 (print edition)

Taxonomy of Peruvian water beetles in the genus Hydrochus Leach, 1817

Key words: Peru, water beetles, Hydrochidae, new species

Materials and methods

Accepted by A. Short: 1 Oct. 2020; published: 23 Oct. 2020 61

HNHM Hungarian Natural History Museum, Budapest, Hungary (G. Makranczy)

Abbreviations for characters on male genitalia figures

adbl: aedeagus dorsal basal lobe

pdmm: paramere dorsal medial margin

Abbreviations used in species descriptions

62 · Zootaxa 4868 (1) © 2020 Magnolia Press PERKINS

Hydrochus alternatus, new species

FIGURE 1. Dorsal habitus of Hydrochus species.

Taxonomy of Peruvian Hydrochus Zootaxa 4868 (1) © 2020 Magnolia Press · 63

64 · Zootaxa 4868 (1) © 2020 Magnolia Press PERKINS

Taxonomy of Peruvian Hydrochus Zootaxa 4868 (1) © 2020 Magnolia Press · 65

Hydrochus astrictus, new species

66 · Zootaxa 4868 (1) © 2020 Magnolia Press PERKINS

Taxonomy of Peruvian Hydrochus Zootaxa 4868 (1) © 2020 Magnolia Press · 67

FIGURE 5. Hydrochus bisinuatus n. sp., habitus and male genitalia of holotype.

68 · Zootaxa 4868 (1) © 2020 Magnolia Press PERKINS

Hydrochus bucapitus, new species

Taxonomy of Peruvian Hydrochus Zootaxa 4868 (1) © 2020 Magnolia Press · 69

FIGURE 6. Hydrochus bucapitus n. sp., habitus and male genitalia of holotype.

70 · Zootaxa 4868 (1) © 2020 Magnolia Press PERKINS

Hydrochus cracentis, new species

FIGURE 7. Hydrochus cracentis n. sp., habitus and male genitalia of holotype.

Taxonomy of Peruvian Hydrochus Zootaxa 4868 (1) © 2020 Magnolia Press · 71

Hydrochus dualis, new species

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FIGURE 8. Hydrochus dualis n. sp., habitus and male genitalia of holotype.

Etymology. Named in reference to the two callosities on the 5th interstria.

Taxonomy of Peruvian Hydrochus Zootaxa 4868 (1) © 2020 Magnolia Press · 73

Hydrochus lescheni, new species

Hydrochus obliquus, new species

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FIGURE 9. Hydrochus lescheni n. sp., habitus and male genitalia of holotype.

Taxonomy of Peruvian Hydrochus Zootaxa 4868 (1) © 2020 Magnolia Press · 75

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Hydrochus parvicollis, new species

Hydrochus spinosus, new species

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Taxonomy of Peruvian Hydrochus Zootaxa 4868 (1) © 2020 Magnolia Press · 79

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Hydrochus tambopaticus, new species

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Hydrochus tripartitus, new species

Hydrochus uniformis, new species

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Taxonomy of Peruvian Hydrochus Zootaxa 4868 (1) © 2020 Magnolia Press · 85

Hydrochus argutus Knisch

Hydrochus argutus Knisch 1921: 3.

Hydrochus bituberculatus Perkins

Hydrochus bituberculatus Perkins, 2019c: 11.

Hydrochus coeneni Makhan

Hydrochus coeneni Makhan 1992: 102

86 · Zootaxa 4868 (1) © 2020 Magnolia Press PERKINS

Hydrochus leei Perkins

Hydrochus leei Perkins, 2019c: 50.