Morphological overview of the amazonicum

Transcription

ZOOTAXA
1274
An interpretation of the morphological variation
in the Simulium amazonicum species group (Diptera: Simuliidae)
of Latin America
A.J. SHELLEY, L.M. HERNÁNDEZ, M.A.MAIA-HERZOG, A.P.A. LUNA DIAS & S.B. LUZ
Magnolia Press
Auckland, New Zealand
A.J. SHELLEY, L.M. HERNÁNDEZ, M.A.MAIA-HERZOG, A.P.A. LUNA DIAS & S.B. LUZ
An interpretation of the morphological variation in the Simulium amazonicum species group
(Diptera: Simuliidae) of Latin America
(Zootaxa 1274)
68 pp.; 30 cm.
27 July 2006
ISBN 978-1-86977-022-8 (paperback)
ISBN 978-1-86977-023-5 (Online edition)
FIRST PUBLISHED IN 2006 BY
Magnolia Press
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ISSN 1175-5326
(Print edition)
ISSN 1175-5334
(Online edition)
Zootaxa 1274: 1–68 (2006)
www.mapress.com/zootaxa/
ISSN 1175-5326 (print edition)
Copyright © 2006 Magnolia Press
ISSN 1175-5334 (online edition)
ZOOTAXA
An interpretation of the morphological variation in the Simulium
amazonicum species group (Diptera: Simuliidae) of Latin America
A.J. SHELLEY1, L.M. HERNÁNDEZ1, M.A.MAIA-HERZOG2, A.P.A. LUNA DIAS2 &
S.B. LUZ3
1
Diptera Division, Department of Entomology, The Natural History Museum, Cromwell Road, London SW7
5BD, UK. E-mail: [email protected]
2
Departamento de Entomologia, Instituto Oswaldo Cruz, Avenida Brasil 4365, Manguinhos, Caixa Postal
926, 21045-900 Rio de Janeiro, R.J., Brasil
3
Centro de Pesquisa Leônidas e Maria Deane, Fundação Oswaldo Cruz, Rua Teresina 476, Adrianópolis,
69.057-070, Manaus, AM, Brazil
Table of contents
Abstract ............................................................................................................................................. 3
Introduction ....................................................................................................................................... 4
Material and methods ........................................................................................................................ 6
Results ............................................................................................................................................... 6
Pupae .......................................................................................................................................... 7
Adults ....................................................................................................................................... 18
Distribution ..................................................................................................................................... 35
Discussion ....................................................................................................................................... 3
Nomenclatural rearrangements involving the amazonicum and siolii groups ................................ 4
Acknowledgements ......................................................................................................................... 45
References ....................................................................................................................................... 4
Appendix ......................................................................................................................................... 49
Abstract
A morphological framework for species within the medically important Simulium amazonicum
group within the subgenus Psaroniocompsa Enderlein is provided as a basis for cytological and
molecular studies. The diagnostic characters for pupae and adults of the group are provided and the
male genitalia are regarded as the major defining feature for the group. Short descriptions of the
pupae and adults of each species are given together with digital images of salient characters, largely
from type specimens. Variation in pupal characters and adult scutal pattern are comprehensively
covered. The taxonomic changes in the amazonicum group from that given in the latest world
inventory of Crosskey and Howard (2004) are as follows: S. argentiscutum Shelley & Luna Dias, S.
Accepted by P. Adler: 13 Jun 2006; published: 27 Jul. 2006
3
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cuneatum (Enderlein), and S. quadristrigatum Enderlein are transferred from “unplaced to group”
to the amazonicum group; S. minusculum Lutz is removed from the amazonicum group to
“unplaced to group”; two new forms (R. Mari and R. Ituxi) are described within S. amazonicum s.l.;
the Argentina, and Manabi forms [previously within S. oyapockense s.l.] are placed together with S.
roraimense s.s. as S. roraimense s.l. The amazonicum group now contains 10 species. Within the
closely related siolii group, S. lourencoi Py-Daniel becomes a junior synonym of S. siolii PyDaniel. The elevation of the siolii group to the new genus Shelleyellum Py-Daniel & Pessoa is not
accepted because of insufficient characters to warrant generic status. Shelleyellum, with its type
species S. siolii, is synonymised with Simulium s.l. Latreille.
Key words: Simuliidae, Neotropical Region, morphotaxonomy, new synonyms, Psaroniocompsa,
Shelleyellum, amazonicum group, siolii group, human onchocerciasis and mansonelliasis vectors
Introduction
In 1905, Goeldi superficially described Simulium amazonicum from human-biting females
collected in the Brazilian Amazon. Work on the distribution and epidemiology of human
mansonelliasis in this region in the 1950s (see Shelley & Coscarón 2001 for review and
literature source) and more recently of human onchocerciasis in the 1970s (see Shelley
2002 for review) provoked several further studies on simuliid taxonomy in the region
because of difficulties in making accurate identifications of newly incriminated vector
species. Initially, S. amazonicum was incriminated as a vector of the parasites responsible
for these two diseases until it was discovered that several species were masquerading
under this name. During this time, it became clear that because of the similarity of females
in some human-biting species, accurate species determination was only possible when
morphological characters in more than one life stage were used. Several species of the
amazonicum group are now known to be vectors. Given the importance of accurate
identification in vector studies on onchocerciasis, the World Health Organization
convened three meetings of simuliid taxonomists (reported in World Health Organization
1979, 1982) to discuss the taxonomy of the Amazonian Simuliidae, with special reference
to the human filariae Mansonella ozzardi (Manson) and Onchocerca volvulus Leuckart.
As a starting point to a revision of S. amazonicum and its relatives, a modern
description of adults, pupae, and larvae of S. amazonicum based on a neotype was
provided by Shelley et al. (1982). In this paper, six other similar species were compared
with S. amazonicum and listed as members of the amazonicum group within the subgenus
Psaroniocompsa. Various papers on S. amazonicum and its relatives were produced at this
stage (e.g. Tidwell et al. 1981b), culminating in two comprehensive taxonomic treatments.
The first, by Py-Daniel (1983), divided the species of the amazonicum group of the
subgenus Psaroniocompsa with a few additions into two new subgenera, Cerqueirellum
and Coscaroniellum [Later Py-Daniel and Moreira Sampaio (1995) elevated these two
subgenera to genera together with subgeneric names of all other Brazilian species]. This
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SHELLEY ET AL.
subgeneric scheme was followed in the revisionary work by Coscarón (1991). A
preliminary revision of the subgenus Psaroniocompsa, with emphasis on the amazonicum
group, was then provided in 1988 by Shelley. A discussion on supraspecific groupings was
given and the subgenera Cerqueirellum and Coscaroniellum were synonymised with
Psaroniocompsa. Most of the species in Cerqueirellum were returned to the amazonicum
group and the rest were unplaced to group; Coscaroniellum species and their synonyms
were placed in the quadrifidum group. Shelley included the following species in the
amazonicum group: S. amazonicum Goeldi, S. chaquense Coscarón, S. oyapockense Floch
and Abonnenc, S. pseudoamazonicum Ramírez Pérez & Peterson, [later synonymised with
S. oyapockense by Shelley et al. (1997)], S. roraimense Nunes de Mello, S. sanguineum
Knab, and S. venezuelense Ramírez Pérez & Peterson. This analysis included discussions
on the inclusion of other names in the group by other authors. Further taxonomic studies
and discussions relevant to the present paper were given by Shelley et al. (1997) on two
amazonicum group species that are vectors of human onchocerciasis in northern Brazil, S.
oyapockense s.l. [referred to as s.l. because it has been used for incorporating synonyms
and is thought to be a species complex] and S. roraimense. In the latest taxonomic and
geographical inventory of world blackflies, Crosskey & Howard (2004) accepted the
species included in the amazonicum group by Shelley (1988) and added S. minusculum
Lutz and S. ganalesense Vargas, Martínez Palácios & Díaz Nájera. A further species group
was added to the subgenus Psaroniocompsa [later regarded as a genus by Py-Daniel &
Moreira Sampaio (1995)] in 1988 by Py-Daniel. He initially published descriptions of
three new species from Brazil within the subgenus Psaroniocompsa: S. damascenoi, S.
lourencoi, and S. siolii. Later (Py-Daniel 1989), he placed these three species in the newly
created siolii group to which he added another new species, S. guaporense. In 2000,
Hamada added a further new species from Brazil to the group, S. tergospinosum. Crosskey
and Howard (2004) recorded these five species within the siolii group.
There is still considerable debate over the supraspecific organisation of the
Neotropical Simuliidae. In this paper, we maintain the species in the amazonicum group as
defined by Shelley (1988) and further assign the following unplaced species in Crosskey
& Howard (2004) to this species group: S. argentiscutum, S. cuneatum, and S.
quadristrigatum.
Taxonomic research on the Simuliidae of Brazil is now moving to a phase of
integrating morphological, cytological, and molecular techniques in order to better
understand morphological variation in relation to species identification and phylogeny.
The amazonicum group is of particular interest because it contains various medically
important species. From our previous work on species of the amazonicum group and the
large number of link-reared specimens accumulated for several of the species, it is now
appropriate to present a morphological framework of S. amazonicum and related species,
taking into account morphological variation in key identification characters. In our current
analysis, more weight is given to the form of the male genitalia than to scutal pattern for
SIMULIUM AMAZONICUM
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the supraspecific arrangement. Our analysis in this paper is based on the pupal and adult
stages because insufficient data are available on larvae of many of the species. It provides
descriptions and images of variations noted in each species on which future cytological
and molecular work can be based.
Material and methods
Our study is a continuation of the work of Shelley et al. (1997), concerning simuliid
species in the Brazilian Amazonia onchocerciasis focus, which placed emphasis on two of
the vector species of human onchocerciasis in Brazil: S. oyapockense s.l. and S.
roraimense, both members of the amazonicum group. Some of the characters already
observed were re-examined in more detail for these two species and in related species, as
well as some characters discussed by Py-Daniel (1983) in order to assess their validity in
identification. Observations were made on link-reared, slide-mounted pupal exuviae
where possible that had been prepared following the method used by Shelley et al. (1997)
so that the characters of the two life stages could be associated. In the pupae, the characters
observed are the length and branching configuration of the gill, the form and distribution
of tubercles on the cephalothorax and frontoclypeus and the number of trichome branches
on these two structures, and the presence or absence of spine combs on the antero-lateral
corners of abdominal tergites II and III. Images of adult scutal patterns were captured
using link-reared specimens, where possible, in two positions: with light anterior and
posterior to the specimens, using the method detailed by Shelley et al. (1997). Tidwell et
al. (1981a,b) made composite drawings of the scutal pattern, with light sources from both
sides, as well as a lateral view of the thorax with either anterior and posterior lighting or
just posterior lighting. We have found our system to be less time consuming and just as
effective. Neither system is perfect because older specimens with damaged setae do not
show perfect scutal patterns, which are dependent on setal form and distribution. Slide
preparations of, in most cases, link-reared adults were used to provide images of the
cibarium and genitalia of the female and the genitalia of the male.
In this paper we have used the term “complex” to refer to a morphologically uniform
species containing cytologically different cytoforms, cytotypes, or cytospecies. We use
sensu lato (s.l.) following a species name to indicate specimens such as topotypes,
cytotypes, and morphoforms together with the primary type(s).
Results
Results are divided into two parts, dealing first with pupae and then with adults. Pupae are
easily categorised by filament number and are grouped accordingly, while adults are dealt
with alphabetically because of the difficulty in grouping scutal patterns.
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SHELLEY ET AL.
PUPAE
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1274
Main diagnostic characters of species within the amazonicum group
Cocoon generally slipper-shaped (Fig. 1), except in S. argentiscutum where it is shoeshaped (Fig. 2). Cocoon light brown, and composed of fibres; margin of anterior aperture
reinforced, weakly to strongly, with or without central protuberance.
Gill white to light brown, consisting of 4–10 forwardly directed filaments branching at
various heights in basal half. Filaments all approximately same length, usually fine (except
S. argentiscutum Fig. 9), edges crenate and distal ends rounded (Fig. 3).
Head with 2+2 frontal and 1+1 facial trichomes that are simple to three branched and
sometimes reduced in length (S. oyapockense s.l.); tubercles absent, rounded, or rounded
and pointed.
Thorax with 5+5 small, simple to 4-branched trichomes; tubercles pointed and/or
rounded with varying distributions. Spine combs absent or present on abdominal tergites II
and III.
Species diagnoses
Table 1 has been constructed by presenting data from the characters being investigated
in groups based on gill filament number. Thus, species with 4 filaments appear at the
beginning of the table followed by those with higher filament numbers and ending with S.
cuneatum, the only species with 10 gill filaments. The pupa of S. quadristrigatum has
never been described. This grouping is to facilitate comparison and has no taxonomic
significance.
Species with four filaments
Only one species within the amazonicum group possesses four gill filaments: S.
venezuelense (Table 1). Gill length is variable, but is short in comparison with other
members of the group. Gill branching is as follows (Fig. 16): the gill trunk divides basally
into two primary branches, each of which then further bifurcates basally. Tubercles are
present on the median portion of the cephalothorax and are rounded in some specimens
and pointed and rounded in others; they are absent from the frontoclypeus (Fig. 6).
Trichomes on the cephalothorax are simple, bifid, or quadrifid; in the frontal region of the
frontoclypeus they are simple and bifid or trifid in the facial region. Spine combs are
absent on tergites II and III of the abdomen.
Species with six filaments
Four species are included in this group: S. argentiscutum, S. ganalesense, S.
oyapockense s.l. (including the synonyms S. cuasisanguineum, S. pseudoamazonicum, and
S. sanchezi and the Bem Querer form [see under ADULTS]) and S. roraimense. Of these,
all are easily distinguishable based on several characters, except S. oyapockense s.l.
[referred to in the broad sense to include populations apart from that of the type locality]
SIMULIUM AMAZONICUM
7
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and S. roraimense, which only differ obviously from one another in the scutal pattern of
the male and possibly in pupal gill length.
FIGURES 1–7. General features of pupae. 1. Slipper-shaped cocoon (S. amazonicum s.s.). 2. Shoeshaped cocoon (S. argentiscutum). 3. Gill filament with crenate edge and rounded tip. 4.
Frontoclypeus with normal trichomes and rounded and pointed tubercles (S. roraimense, topotype).
5. Frontoclypeus with reduced trichomes and rounded tubercles (S. oyapockense, topotype). 6.
Frontoclypeus without tubercles (S. venezuelense). 7. Spine combs on abdominal tergite III (S.
oyapockense, topotype).
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SHELLEY ET AL.
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1274
FIGURES 8–10. Respiratory gill of pupa. 8. S. amazonicum, neotype. 9. S. argentiscutum,
paratype. 10. S. chaquense.
SIMULIUM AMAZONICUM
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FIGURES 11–13. Respiratory gill of pupa. 11. S. cuneatum (after Coscarón, 1991). 12. S.
ganalesense, paratype. 13. S. oyapockense, topotype.
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SHELLEY ET AL.
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1274
FIGURES 14–16. Respiratory gill of pupa. 14. S. roraimense, topotype. 15. S. sanguineum. 16. S.
venezuelense.
SIMULIUM AMAZONICUM
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FIGURES 17–22. Variations in respiratory gill configuration. 17. S. amazonicum, topotype, R.
Purus. 18. Left hand gill of specimen in Fig. 17. 19. S. amazonicum, topotype, R. Purus. 20. S.
amazonicum, topotype, R. Purus. 21. S. amazonicum s.l., R. Ituxi. 22. S. amazonicum s.l., R. Ituxi.
12
SHELLEY ET AL.
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1274
FIGURES 23–26. Variations in respiratory gill configuration. 23. S. ganalesense, Belize form. 24.
S. ganalesense, Belize form. 25. S. oyapockense s.l., Bem Querer. 26. S. oyapockense s.l., R. Ituxi.
Gill length (average) in these species varies from 0.8–1.8 mm, with S. argentiscutum
having the shortest (0.8 mm) and the other three the longest (1.8 mm). Large ranges in
length were seen (Table 2) in the following species: S. ganalesense (type locality—1.6–2.1
mm; Belize—2.4–3.9 mm), S. oyapockense s.l. (type locality—0.6–2.1 mm; other
localities—0.8–1.9 mm), S. roraimense (type locality—1.6–2.6 mm; other localities—
1.4–2.0 mm).
The gill trunk initially divides basally into three primary branches. The dorsal, median
and ventral branches then bifurcate at different distances from the gill base depending on
species. The broad form of the gill filaments serves to distinguish S. argentiscutum (Fig. 9)
from the other species (Figs. 12–14) where the gill is more filamentous. Identification of
the remaining three species is more difficult, being based on different branching
configurations, which themselves vary. The more distal branching of the dorsal and
median bifurcations in comparison to the ventral in S. ganalesense Belize form (Figs. 23,
24) distinguishes this species from S. ganalesense from its type locality in Mexico (Fig.
SIMULIUM AMAZONICUM
13
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12) and all the other species. The branching configuration of S. oyapockense s.l. was
described by Shelley et al. (1997) (Figs. 13, 25, 26), and found to be similar in specimens
of its synonyms S. cuasisanguineum, S. pseudoamazonicum, and S. sanchezi as well as to
S. roraimense (Fig 14). Typically, the dorsal bifurcation is most basal and the median and
ventral bifurcations, at the same level, slightly more distal to it (Figs. 13, 14). Common
variations are with the ventral slightly more distal to the median (Fig. 25) or with both
median and ventral branches at the same level, but significantly more distal from the base
than the dorsal branch (Fig. 26). Exceptionally, the ventral bifurcation in S. oyapockense
s.l. can be significantly more distal, and in S. roraimense the median bifurcation can be the
most distal.
Tubercle form may be useful for distinguishing some of the species. Tubercles are
found on the anterior and median portions of the cephalothorax in all species, except S.
argentiscutum where they also occur on the posterior portion, and S. cuasisanguineum
(regarded as a synonym of S. oyapockense s.l.) on only the median portion. In all species
the tubercles are both pointed and rounded, except in S. argentiscutum where only rounded
are present. The density of tubercles on both the cephalothorax and frontoclypeus is
variable within populations of the same species. On the frontoclypeus, only S.
cuasisanguineum (a synonym of S. oyapockense s.l.) is devoid of tubercles (as in S.
venezuelense, Fig. 6). In all the other species in the “six filament group” only rounded
tubercles are present on the frontoclypeus (as in S. oyapockense s.s., Fig. 5), except in S.
roraimense (Fig. 4) and the sanchezi form of S. oyapockense s.l. [only a small number was
examined] where pointed ones are also present. However, some exceptions occur where
male scutal pattern indicates either S. oyapockense s.l. or S. roraimense, but which show a
tubercle form not typically seen in most populations of these distinct species. Thus,
pointed tubercles are present in males at the rivers Jacy Paraná, Ituxi, Toototobi, and
Arraia in Brazil that have a scutal pattern resembling that of S. oyapockense s.l. Also under
this category are specimens from Argentina, Ecuador, and Paraguay. In the case of the
material from Ecuador, the male scutal pattern is of the S. roraimense type, but the
tubercles are rounded as in typical S. oyapockense s.l. The two females from Paraguay
show pointed and rounded tubercles, but in the absence of males could not be identified to
species.
Trichome branching provides no means of species separation, but the reduced
trichome length in S. oyapockense s.l. separates this species from the rest (Table 1; Figs. 4,
5). However, there are some exceptions. In specimens from the R. Ituxi and Jacy Parana in
Brazil, reduced trichomes were not obvious despite the oyapockense-type male scutal
pattern. None of the specimens from Paraguay had reduced trichomes. However, this
character is difficult to see and to assess without access to several of the species for
comparison and hence can be done at only museum level and not in the field. Spine combs
are absent on tergites II and III in all species except S. oyapockense s.l. (Fig. 7) and one of
its synonyms, S. sanchezi. However, spine combs are absent on one or both of the tergites
in some specimens in populations of S. oyapockense s.l. (Table 1).
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SHELLEY ET AL.
ZOOTAXA
TABLE 1. Morphological characters of pupae of S. amazonicum group species.
Species
Locality
Cephalothorax
Frontoclypeus
Number
Gill Gill length
examined filament in mm.
number Mean, S.D.,
Range (no. Tubercles Trichome Tubercles Trichomes
exam.)
branches
Facial Frontal
S. venezuelense
&
4
1.3 (1)
Mr
1,2
none
2
1
n.d.
%
4
1.4 (1)
Mpor
1,2
none
2
1
-, - (1)
%
4
n.d.
Mr
1,2
none
2
1
n.d.
&&, 3%%
4
0.9-1.2 (3)
Mpor
1,2,4
none
2,3
1
-, - (2)
&&, 2%%
6
0.8-0.9 (3)
AMPr
1,2
r
1,2
1,2
-, - (3)
1 ,1
& %
6
2.4-3.9**
AMpor
1,2
r
2
1,2
-,- (2)
&&, 1%
6
AMpor
1,2,3
r
1,2
1 (r)
+, + (2)
&&, 8%%
1.84; 0.2;
1.6-2.1 (5)
6
1,2,3
r
1,2
1 (r)
&&, 2%%
1.1; 0.16; AMpor
0.8-1.3 (14)
+, + (8);
-, +(1)
6
0.9; 0.06;
0.9-1.0 (4)
AMpor
1,2
r
1,2
1 (r)
+, + (1);
-, + (2)
Amapá, Cach.
Patanari
1
Amapá, Cach.
Patanari
1
Amapá, R.
Patanari
1
Cach. 2
S. argentiscutum
Cach. Teotonio
3
2
S. ganalesense
Belize
S. oyapockense s.l.
R. Oyapock
(type loc.)
Spine
combs
on tergites II,
III (no.
exam.)
8
C. Bem
Querer
7
R. Aila
2
R. Ajarani 1
4
%%
6
1,2
r
2
1,2
-, - (3)
R. Arraia
5
&&
1.5; 0.2; AMPpor
1.2-1.9 (4)
6
AMpor
1,2
r
1
1 (r)
+, + (4)
R. Cauamé
3
%%
1.3; 0.09;
1.2-1.4 (4)
6
0.9-1.3 (2)
AMpor
1,2
r
1,2
1 (r)
n.d.
&&, 2%%
6
1,2
por
1,2
1,2
-, - (6)
&&, 3%%
1.6; 0.1; AMPpor
1.5-1.9 (5)
6
1,2
por
1,2
1
-, - (1)
%%
2.0; 0.2; AMPpor
1.9-2.5 (5)
6
1.6-1.9 (2) AMPpor
1,2
r
1
1,2
-, - (1)
R. Ituxi
4
R. Jacy Paraná 2
R. Madeira, C.
Teotonio
2
................to be continued
SIMULIUM AMAZONICUM
15
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ZOOTAXA
TABLE 1 (continued)
1274
Species
Locality
Cephalothorax
Frontoclypeus
Number
Gill Gill length
number Mean, S.D.,
exam.)
branches
Facial Frontal
Spine
combs
on tergites II,
III (no.
exam.)
&
6
1.0 (1)
AMpor
1,2
r
1,2
1 (r)
n.d.
R. Maú, Gua- 2
nabara
&&, 4%%
6
AMpor
1,2
r
1,2
1 (r)
+, + (3);
-, -(1)
R. Maú,
Uiramutã
&&, 6%%
1.0; 0.1;
0.9-1.2 (4)
6
1.2; 0.1;
1.1-1.4 (6)
AMpor
1,2
r
1,2
1 (r)
+, + (7)
6
1.7 (1)
AMpor
1,2
r
1,2
1 (r)
n.d.
6
1.3 (1)
AMpor
1,2
r
1,2
1 (r)
+, + (1)
& %%
6
AMpor
1,2
r
1,2
1 (r)
%%
1.6; 0.2;
1.2-1.9 (5)
+, + (4);
-, + (1)
6
1.3-1.4 (2)
AMpor
1,2,4
r
2
2 (r)
-, + (1)
& %
6
2.1 (1)
AMPpor
2
r
2
1
-, - (2)
&&, 4%%
6
AMpor
1,2
r
1,2
1 (r)
&&, 1%
1.0; 0.3;
0.8-1.2 (7)
+, + (2);
-, + (1)
6
-
AMpor
1,2
r
1,2
1 (r)
&&
+, + (1);
-, + (1)
6
1.7 (2)
AMPpor
2
por
n.d.
1,2
n.d.
%
6
1.7 (1)
Mpor
1,2,3,4
None
2
1
-, - (1).
&&, 3%%
6
1.4-1.6 (2) AMPpor
1,2,4
r
1,2
1
-, - (1)
& %
6
1.2-1.3 (2) AMPpor
1,2,3
por
2
1,2
+, + (1)
&&, 7%%
6
1,2
por
1,2
1
-, - (10)
%
1.8; 0.2; AMPpor
1.6-2.6 (11)
6
1,2
por
1,2
1
-, - (1)
1,2,3
por
1,2
1
-, - (6)
R. Maú, Cariri
1
3
&
1%
R. Mucajai
1
R. Murupu
R. Tacutu,
Lethem
1 ,5
R. Tacutu, C.
M. de P.
2
R. Toototobi
1 ,1
R. Uraricoeira 3
R. Urubu
2
Paraguay
2
Synonyms of
S. oyapockense s.l.
S. cuasisanguineum
1
S. pseudoam- 3
azonicum
S. sanchezi
S. roraimense
R. Cauamé
(type loc.)
R. Ajarani 1
R. Arraia
1 ,1
8
1
&&,
%%
10
6
6
n.d.
AMPpor
1.7; 0.1; AMPpor
1.4-2.0 (12)
................to be continued
16
SHELLEY ET AL.
ZOOTAXA
TABLE 1 (continued)
Species
Locality
R. Catrimani
Cephalothorax
Frontoclypeus
Number
Gill Gill length
number Mean, S.D.,
exam.)
branches
Facial Frontal
&&, 9%%
6
%
6
1.6 (1)
&&, 5%%
6
1.5; 0.21;
1.2-1.8
9
R.Maú, Cariri
Ecuador, R.
Portoviejo
1
4
S. amazonicum
&&,
%%
R.Purus
por (12)
r(1)
1,2
1
-, - (4)
AMPpor
1,2
por
2
1
-, - (1)
AMPpor
1,2
r
1,2
1,2
-,-(8)
1.4; 0.18; AMPpor
1.0-1.8 (20)
1,2,3,4
por
2,3
1,2,3
-, - (14)
&
8
1.8-2.0*** AMPpor
2
por
1
1
-, - (1)
&
8
AMPpor
2
por
1
1
-, - (1)
&&, 4%%
8
1.6; 0.05; AMPpor
1.5-1.7 (5)
1,2,3
r
1,2
1
-, - (7)
?
10
1
por
1
1
?
Arroyo Carolis
1
Arroyo
Zapirain
1
R.Tagachi
1,2
8
10
10
S. chaquense
S. sanguineum
1.6; 0.1; AMPpor
1.5-1.7 (10)
Spine
combs
on tergites II,
III (no.
exam.)
4
S. cuneatum
****
Argentina,
Corrientes
1.2-1.8
por
Tubercle distribution and form on cephalothorax and frontoclypeus
A—anterior part; M—median part; P—posterior part—symbol underlined shows area of most
density
po—pointed tubercles ; r—rounded tubercles
Trichomes
(r)—Reduced trichomes; 1—simple; 2—bifid; 3—trifid; 4—quadrifid
n.d.—No data
* Based on Wygodzinsky (1961). **Measurements of Belize form. Paratypes of S. ganalesense 1.8,
0.35, 1.6-2.1 (8). *** Based on Coscarón (1971) ****Based on Coscarón (1983).
SIMULIUM AMAZONICUM
17
1274
ZOOTAXA
1274
Species with eight filaments
The following three species occur in this group: S. amazonicum, S. chaquense, and S.
sanguineum.
The overlap in gill length between species precludes the use of this character for
identification (Table 1), but configuration of gill branching may be used (Figs. 8, 10, 15).
The gill branches basally into three primary branches. The dorsal and median primary
branches then each further divide into three secondary filaments and the ventral primary
branch into two secondary filaments. Distance of branching of the secondary filaments
from the gill base varies with species. Despite the variation in gill branching in S.
amazonicum (Figs. 17–22), often varying between the left hand side and right hand side of
the same specimen, this species may be easily distinguished from S. sanguineum (Fig. 15).
In the latter species, bifurcations on the dorsal and median primary branches are more
distal than those in S. amazonicum and also the most distal bifurcations of the secondary
dorsal and median branches of the gill are on the lower part in S. sanguineum and on the
upper part in S. amazonicum. Variation in gill branching height has been observed in S.
sanguineum by Tidwell et al. (1981a). Simulium chaquense (Fig. 10) is insufficiently
distinct to separate it from S. amazonicum. Both pointed and rounded tubercles are present
on all portions of the cephalothorax of all three species. On the frontoclypeus, only S.
sanguineum has rounded tubercles, whereas both pointed and rounded occur in the other
two species. Trichome branching and length do not show any difference between the three
species and all lack spine combs on tergites II and III. Py-Daniel (1983) referred to the
presence of spine combs on tergites II and IV of S. chaquense, but none was found on the
two specimens examined by us.
Species with ten filaments
Only S. cuneatum occurs in this group and few data are available on its pupal
morphology. Its pupal gill with 10 filaments is sufficient to distinguish it from the other
species of the amazonicum group. Gill branching is basal with three primary filaments
dividing (from dorsal to ventral) into four, four and two secondary filaments respectively
(Fig. 11).
ADULTS
This section on adult flies first provides brief descriptions and images of characters that
define the species of the amazonicum group. It then deals wholly with the adult scutal
pattern, which is the only adult character that significantly varies at the population level. In
species that are morphologically close in scutal pattern, links are made to the pupal stage
where link-reared material is available.
18
SHELLEY ET AL.
Main diagnostic characters of species within the amazonicum group
Females
Nudiocular area absent in all species, except S. venezuelense where it is highly
reduced (Figs. 27–36). Cibarium with slightly developed, sclerotised cornuae and wellsclerotised margin to central trough; armed with several rows of sharp teeth along margin
of central trough, which are absent from small median area (Figs. 37–46).
Scutum black and silvery grey pruinose with a distinct pattern and numerous,
recumbent, silvery grey hairs interspersed with semi-erect black hairs. The scutal patterns
illustrated (Figs. 47–66) are based on type specimens where possible. Variation in this
pattern is dealt with below. The scutal pattern in each species is a variation on a basic
scheme. This consists of a silvery grey pruinose scutum with a black median vitta running
from anterior scutal border to about four-fifths of scutal length that broadens posteriorly,
1+1 lateral black vittae and 1+1 intervittal bands, which are black or silvery grey
depending on light direction. This pattern varies with light direction, so for each species
the first image has been produced when the light source is anterior to the specimen and the
second image when the light source is posterior.
Eighth sternite with sclerotised posterior margins with groups of setae; gonapophyses
unsclerotised, sub-triangular, membranous and covered in fine setae (Figs. 91–100). Cerci
hemispherical, paraprocts poorly developed and sub-rectangular and sclerotised to varying
extent (Figs. 101–110). Genital fork (Figs. 111–120) slightly sclerotised with poorly
developed lateral arms and anterior processes.
Males
Scutum velvet black with silvery grey pruinosity forming pattern. Basic pattern for all
species of 1+1 sub-median, silvery grey cunae (triangles) of various sizes on anterior
border of scutum and silvery grey lateral and posterior scutal margins (Figs. 121–144).
Gonocoxite slightly longer than wide; gonostyle slender, sub-triangular, about fourfifths length of gonocoxite and with one sub-terminal spine (Figs. 145–153) [Coscarón
(1971) records 2 spines in S. chaquense, while only 1 was observed in our material.].
Ventral plate very lightly sclerotised, crescent-shaped with many fine setae and with
median part of anterior margin bluntly pointed and basal arms poorly developed (Figs.
154–162). Paramere sclerotised, with well-developed spines (Figs. 163–171) [the lightly
sclerotised paramere of S. ganalesense is due to 60 years preservation of specimen in
alcohol causing breakdown of sclerotin.]. The male of S. quadristrigatum is not known.
Species diagnosis using scutal patterns
A short description of the two scutal patterns dependent on light source follows for
each species and each sex. Also affecting the coloration of the scutal pattern is the
condition of the specimen, especially in older type material. The following descriptions
are mostly based on link-reared specimens. The similarity of scutal patterns makes
accurate identification impossible without recourse to pupal morphology.
SIMULIUM AMAZONICUM
19
ZOOTAXA
1274
ZOOTAXA
1274
Females
Simulium amazonicum neotype (Figs. 47, 48) from 8-filamented pupa—scutum
largely silvery grey pruinose with broad median vitta, reduced lateral vittae and very
indistinct intervittal bands irrespective of light direction.
Two variations have been recorded at the R. Ituxi and the R. Mari, both tributaries of
the R. Purus and close to the type locality. In the R. Ituxi form (Figs. 67, 68), black
intervittal bands are present (anterior light source) that greatly resemble variations seen in
S. oyapockense s.l. from both Brazil and neighbouring countries (Figs. 57, 58, 71–88).
Simulium amazonicum Ituxi form may be distinguished by the lateral vittae not reaching
the anterior scutal margin with a posterior light source, whereas these vittae do in S.
oyapockense s.l. In the R. Mari form of S. amazonicum (Figs. 69, 70) the black intervittal
bands are highly developed (anterior light source) and the lateral vittae (posterior light
source) reach the anterior scutal border as in S. oyapockense s.l. The only means of
distinguishing this form is in reared specimens by the 8-filamented gill compared to the 6filamented gill in S. oyapockense s.l.
Simulium argentiscutum paratype (Figs. 49, 50) from 6-filamented pupa—scutum
largely silvery grey pruinose and median vitta very fine; with light source anterior lateral
vittae broader than median and run for almost whole length of scutum and intervittal bands
broad, crescent-shaped and running from anterior border to about the mid part of scutum;
with light source posterior lateral vittae thin and shorter and intervittal bands faintly
visible as silvery commas.
Simulium chaquense paratype (Figs. 51, 52) from 8-filamented pupa—with anterior
light source median vitta relatively narrow, lateral vittae relatively broad, but not reaching
anterior scutal border, and intervittal bands well developed in form of commas extending
from anterior scutal border to mid point of scutum; with posterior light source median vitta
broader, lateral vittae broader and extending to anterior scutal border, intervittal bands
indistinct silvery commas.
Simulium cuneatum (Figs. 53, 54) from 10-filamented pupa—with similar pattern to S.
chaquense, except median vitta broad irrespective of light direction and intervittal bands,
seen with anterior light source, slightly shorter.
Simulium ganalesense paratype (Figs. 55, 56) with 6-filamented pupa—median vitta
broad irrespective of light source, lateral vittae broad and well developed and only
extending to scutal border with posterior light source; intervittal bands large and commashaped with anterior light, but less distinct and silver with posterior light source.
Simulium oyapockense topotype (Figs. 57, 58) with 6-filamented pupa, S. roraimense
topotype (Figs 61, 62) with 6-filamented pupa and S. sanguineum holotype (Figs. 63, 64)
with 8-filamented pupa—almost indistinguishable in scutal pattern from S. ganalesense
with 6-filamented pupa.
20
SHELLEY ET AL.
ZOOTAXA
1274
FIGURES 27–36. Female nudiocular area. 27. S. amazonicum, topotype. 28. S. argentiscutum,
paratype. 29. S. chaquense, paratype. 30. S. cuneatum. 31. S. ganalesense, paratype. 32. S.
oyapockense, topotype. 33. S. quadristrigatum, holotype. 34. S. roraimense, topotype. 35. S.
sanguineum, paratype. 36. S. venezuelense.
SIMULIUM AMAZONICUM
21
ZOOTAXA
1274
FIGURES 37–46. Female cibarium. 37. S. amazonicum topotype. 38. S. argentiscutum, paratype.
39. S. chaquense, paratype. 40. S. cuneatum. 41. S. ganalesense, paratype. 42. S. oyapockense,
topotype. 43. S. quadristrigatum, holotype. 44. S. roraimense, topotype. 45. S. sanguineum,
paratype. 46. S. venezuelense.
We record variation in S. oyapockense s.l. with its widespread distribution in Brazil
and in Colombia, Ecuador and Uruguay. Minor variations occur in the shape of the vittae,
but more obvious differences occur in the form of the intervittal cunae (Figs. 57, 58,
71–88). Similarly, slight variation in scutal pattern occurs in S. roraimense s.l. from E
cuador (Figs. 89, 90).
Simulium venezuelense (Figs. 65, 66) with 4-filamented pupa—irrespective of light
source with median and lateral vittae broad, latter reaching anterior scutal border and
intervittal bands comma-shaped and indistinct.
22
SHELLEY ET AL.
ZOOTAXA
1274
FIGURES 47–54. Female scutal pattern. The first figure for each species shows anterior light
direction and the second posterior light direction. 47, 48. S. amazonicum, neotype. 49, 50. S.
argentiscutum, paratype. 51, 52. S. chaquense, paratype. 53, 54. S. cuneatum.
Males
Simulium amazonicum topotype (Figs. 121, 122) from 8-filamented pupa—1+1 submedian, silver triangles well developed, extending for half of scutal length with anterior
light source; triangles very indistinct with posterior light source.
Simulium argentiscutum paratype (Figs. 123, 124) from 6-filamented pupa—as in S.
amazonicum, but posterior tip of cunae curved.
Simulium chaquense (Figs. 125, 126) from 8-filamented pupa—scutal pattern well
developed as in S. amazonicum, but with posterior part of cunae almost extending to
posterior silver border of scutum and 1+1 ovoid black areas in anterior part of cunae with
light anterior; scutal pattern still evident but less distinct with posterior light source.
Simulium cuneatum holotype (Figs. 127, 128) [species subsequently redescribed and
associated with 10- filamented pupa by Coscarón (1983)]—scutal pattern similar to that of
S. chaquense with both light directions.
Simulium ganalesense paratype (Figs. 129, 130) from 6-filamented pupa—silver
cunae large and extending to join posterior silver pruinose margin of scutum with anterior
light; similar pattern with posterior light source, except cunae connect anteriorly to lateral
border of scutum.
SIMULIUM AMAZONICUM
23
ZOOTAXA
1274
FIGURES 55–70. Female scutal pattern. The first figure for each species shows anterior light
direction and the second posterior light direction. 55, 56. S. ganalesense, paratype. 57, 58. S.
oyapockense, topotype. 59, 60. S. quadristrigatum, holotype. 61, 62. S. roraimense, topotype. 63,
64. S. sanguineum, holotype. 65, 66. S. venezuelense. 67–70. Variations in female scutal pattern, S.
amazonicum s.l. 67, 68. R. Ituxi. 69, 70. R. Mari.
24
SHELLEY ET AL.
ZOOTAXA
1274
FIGURES 71–86. Variations in female scutal pattern. S. oyapockense s.l. — Brazil, localities north
of R. Amazon. 71, 72. Cachoeira Bem Querer. 73, 74. R. Tacutu. 75, 76. R. Pium. 77, 78. R. Mau. S.
oyapockense s.l. — Brazil, localities south of R. Amazon. 79, 80. R. Ituxi. 81, 82. R. Jacy Paraná.
83, 84. R. Araguaia. 85, 86. Countries other than Brazil: Colombia, Caño Cuduyari.
SIMULIUM AMAZONICUM
25
ZOOTAXA
1274
FIGURES 87–90. Variations in female scutal pattern. S. oyapockense s.l. Countries other than
Brazil. 87, 88. Uruguay, R. Dayman. S. roraimense Ecuador form. 89, 90. Ecuador, R. Portoviejo.
In the Belize form of S. ganalesense (Figs. 131, 132), the scutal pattern is close to that
of S. argentiscutum, S. chaquense, and S. cuneatum in that the cunae extend more
posteriorly and curve at the tips and contain ovoid black areas with anterior lighting; this
pattern is apparent, but indistinct, with posterior lighting.
Simulium oyapockense topotype (Figs. 133, 134) from 6-filamented pupa—scutal
pattern very similar to that of S. amazonicum.
Simulium roraimense holotype and topotype (Figs. 135–138) from 6-filamented
pupa—silver cunae extend posteriorly to join posterior pruinose margin of scutum and 1+1
small, black, sub-median cunae on anterior margin of scutum within these silver cunae
with anterior lighting; with posterior lighting silver cunae thinner and their contained
black cunae less apparent. [Because of the poor quality of the image of the holotype, an
image of a topotype has also been provided.]
Two morphoforms occur. In Ecuador, males from the R. Portoviejo (Manabi form)
have the sub-median black vitta visible with an anterior light source replaced with a
roughly square-shaped black mark (Fig. 139), that is unclear with posterior lighting (Fig.
140). The Argentina form from several localities in this country (Map 2) is similar to S.
roraimense topotypes, with minor differences in male scutal pattern (see Coscarón &
Coscarón-Arias 2000). We have placed it as a separate form because of the distance of the
localities in Argentina to the type locality and the restricted distribution of S. roraimense
s.s. to Roraima State, Brazil.
Simulium sanguineum (Figs. 141, 142) from 8-filamented pupa—silver cunae reach
beyond mid point of scutum. Tidwell et al. (1981a) recorded variation in the length of the
tails of the silver cunae, which sometimes join with the silver pruinose posterior scutal
margin. Scutal pattern indistinct with posterior lighting.
Simulium venezuelense (Figs. 143, 144) from 4-filamented pupa—silver cunae
expanded laterally to pruinose lateral margin of scutum appearing as sub-ovoid areas with
anterior lighting; scutum black with posterior lighting.
26
SHELLEY ET AL.
ZOOTAXA
1274
FIGURES 91–100. Eighth sternite and gonapophyses. 91. S. amazonicum, topotype. 92. S.
argentiscutum, topotype. 93. S. chaquense, paratype. 94. S. cuneatum. 95. S. ganalesense, paratype.
96. S. oyapockense, topotype. 97. S. quadristrigatum, holotype. 98. S. roraimense, topotype. 99. S.
sanguineum. 100. S. venezuelense.
SIMULIUM AMAZONICUM
27
ZOOTAXA
1274
FIGURES 101–110. Cercus and paraproct. 101. S. amazonicum, topotype. 102. S. argentiscutum,
topotype. 103. S. chaquense, paratype. 104. S. cuneatum. 105. S. ganalesense, paratype. 106. S.
oyapockense, topotype. 107. S. quadristrigatum, holotype. 108. S. roraimense, topotype. 109. S.
sanguineum. 110. S. venezuelense.
28
SHELLEY ET AL.
ZOOTAXA
1274
FIGURES 111–120. Genital fork. 111. S. amazonicum topotype. 112. S. argentiscutum, topotype.
113. S. chaquense, paratype. 114. S. cuneatum. 115. S. ganalesense, paratype. 116. S. oyapockense,
topotype. 117. S. quadristrigatum, holotype. 118. S. roraimense, topotype. 119. S. sanguineum. 120.
S. venezuelense.
SIMULIUM AMAZONICUM
29
ZOOTAXA
1274
FIGURES 121–136. Male scutal pattern. The first figure for each species shows anterior light
direction and the second posterior light direction. 121, 122. S. amazonicum, topotype. 123, 124. S.
argentiscutum, paratype. 125, 126. S. chaquense. 127, 128. S. cuneatum, holotype. 129, 130. S.
ganalesense, paratype. 131, 132. S. ganalesense, Belize form. 133, 134. S. oyapockense, topotype.
135, 136. S. roraimense, holotype.
30
SHELLEY ET AL.
ZOOTAXA
1274
FIGURES 137–144. Male scutal pattern. The first figure for each species shows anterior light
direction and the second posterior light direction. 137, 138. S. roraimense, topotype. 139, 140. S.
roraimense Ecuador form, Ecuador, R. Portoviejo. 141, 142. S. sanguineum topotype. 143, 144. S.
venezuelense.
As a result of the above findings, our list of species for the amazonicum group is as
follows:
amazonicum Goeldi, 1905.
tallaferoae Ramírez-Pérez, 1971.
R. Mari form’ [From this paper.]
‘R. Ituxi form’ [From this paper.]
argentiscutum Shelley & Luna Dias, 1980.
‘E’ of Tidwell, Peterson, Ramírez Pérez, Tidwell & Lacey, 1981b.
‘Madeira’ of Shelley, Pinger & Moraes, 1982.
chaquense Coscarón, 1971.
cuneatum (Enderlein, 1936).
ganalesense Vargas, Martínez Palácios & Díaz Nájera, 1946.
‘Belize form’ of Shelley, Hernández & Penn, 2002.
oyapockense Floch & Abonnenc, 1946. [Complex.]
pseudosanguineum Ramírez Pérez & Peterson, 1981.
pseudoamazonicum Ramírez Pérez & Peterson, 1981.
cuasisanguineum Ramírez Pérez, Yarzábal & Peterson, 1982.
sanchezi Ramírez Pérez, Yarzábal & Peterson, 1982.
SIMULIUM AMAZONICUM
31
ZOOTAXA
1274
FIGURES 145–153. Male gonocoxite and gonostyle. 145. S. amazonicum, topotype. 146. S.
argentiscutum, paratype. 147. S. chaquense. 148. S. cuneatum, holotype. 149. S. ganalesense,
holotype. 150. S. oyapockense, topotype. 151. S. roraimense, topotype. 152. S. sanguineum. 153. S.
venezuelense.
32
SHELLEY ET AL.
ZOOTAXA
1274
FIGURES 154–162. Male ventral plate. 154. S. amazonicum, topotype. 155. S. argentiscutum,
paratype. 156. S. chaquense. 157. S. cuneatum, holotype. 158. S. ganalesense, paratype. 159. S.
oyapockense, topotype. 160. S. roraimense, topotype. 161. S. sanguineum. 162. S. venezuelense.
‘Bem Querer’ form of Procunier, 1988 [as cytoform of S. oyapockense s.l.].
‘A’, ‘B’, ‘C’ of Tidwell, Peterson, Ramírez-Pérez, Tidwell & Lacey, 1981b.
‘Barbacoas’ of Shelley, Pinger & Moraes, 1982.
quadristrigatum Enderlein, 1934.
roraimense Nunes de Mello, 1974.
‘Argentina form’ [as S. oyapockense s.l. of Coscarón & Coscarón-Arias, 2000] [From
this paper.]
‘Manabi form’ Procunier, Shelley & Arzube 1987 [as cytoform of S. oyapockense s.l.]
[From this paper.]
sanguineum Knab, 1915.
venezuelense Ramírez Pérez & Peterson, 1981.
SIMULIUM AMAZONICUM
33
ZOOTAXA
Source references for descriptions of species
1274
amazonicum Goeldi, 1905. (Py-Daniel 1983; Shelley et al. 1982).
argentiscutum Shelley & Luna Dias, 1980. (Original authors; Py-Daniel 1983).
chaquense Coscarón, 1971. (Original author; Coscarón 1983; Py-Daniel 1983).
cuneatum (Enderlein, 1936). (Coscarón 1983; Py-Daniel 1983).
ganalesense Vargas, Martínez Palácios & Díaz Nájera, 1946. (Original authors; Shelley et
al. 2002).
oyapockense Floch & Abonnenc, 1946. (Py-Daniel 1983; Shelley et al. 1997).
quadristrigatum Enderlein, 1934. (Coscarón 1983).
roraimense Nunes de Mello, 1974. (Py-Daniel 1983; Shelley et al. 1997).
sanguineum Knab, 1915. (Tidwell et al. 1981a).
venezuelense Ramírez Pérez & Peterson, 1981. (Original authors; Py-Daniel 1983).
FIGURES 163–171. Male paramere. 163. S. amazonicum, topotype. 164. S. argentiscutum,
paratype. 165. S. chaquense. 166. S. cuneatum, holotype. 167. S. ganalesense, holotype. 168. S.
oyapockense, topotype. 169. S. roraimense, topotype. 170. S. sanguineum. 171. S. venezuelense.
34
SHELLEY ET AL.
Distribution
ZOOTAXA
1274
Vectors of M. ozzardi and O. volvulus in Brazil
In Map 1, we show the distributions of four vector species in Brazil that have been
relatively well studied. These are S. amazonicum s.l. and S. argentiscutum (vectors of
Mansonella ozzardi), S. oyapockense s.l. and S. roraimense s.l. (vectors of O. volvulus).
MAP 1. The distribution of S. amazonicum s.l., S. argentiscutum, S. oyapockense s.l., and S.
roraimense s.l. of the amazonicum group in Brazil.
SIMULIUM AMAZONICUM
35
ZOOTAXA
1274
Simulium amazonicum s.l. is restricted to several southern tributaries of the R.
Amazon and one locality in Roraima State (Material Examined). The southern tributaries
cover a vast and geologically similar area of the Amazon basin that has been poorly
prospected for simuliids because of its difficult access. Outside Brazil (Map 2), this
species has been recorded in the Amazonian region of Colombia (Tidwell et al. 1981b),
and from the Upper Orinoco region adjacent to Brazilian Amazonia and the north of
Venezuela (Ramírez Pérez 1983).
MAP 2. The distribution of S. amazonicum s.l., S. argentiscutum, S. oyapockense s.l., and S.
roraimense s.l. of the amazonicum group in the Neotropical Region.
36
SHELLEY ET AL.
Simulium argentiscutum is restricted to the upper Amazon region of Brazil (R.
Solimões) and the adjacent R. Amazonas (Colombia) and some southern tributaries of the
Amazon in Brazil (Maps 1 and 2) (Material Examined; Py-Daniel 1983). It also occurs
westwards in Peru and with more intensive prospection will no doubt be found in the
intervening areas of western Brazilian Amazonas state. A single record exists from Bahia
State in Brazil.
Simulium oyapockense s.l. has a wide distribution within Brazil from northern
Roraima to the south of the country (Map 1). It is more commonly found breeding in
northern Amazon tributaries and is widespread here. It appears to occur more infrequently
in southern Amazon tributaries, but more prospection is needed in this vast area to
establish its true distribution. It also occurs, but less frequently, in southern Brazil where
simuliid prospection has been more concentrated and in western Brazil, but more
prospection is needed to get a clearer picture of its distribution here. It is also recorded
(Map 2) from Colombia (Tidwell et al. 1981b), French Guiana (Floch & Abonnenc 1946;
Hamada & Fouque 2001), and Guyana (Material Examined; Shelley et al. 2004) and is
almost certainly present in Surinam. In Venezuela, it occurs in the Orinoco headwaters and
to the north and east of the country (Ramírez Pérez 1983; Ramírez Pérez et al. 1982) [we
have included the synonyms S. cuasisanguineum, S. pseudoamazonicum, S.
pseudosanguineum, and S. sanchezi in the maps under S. oyapockense s.l.]. Human-biting
females collected in Bolivia (Material Examined) are also probably of this species, and it
undoubtedly occurs in Peru in the lowland rain forest region adjacent to Amazonas and
Rondônia states of Brazil.
Simulium roraimense s.s. has been recorded from only a few localities in the savannas
and forests of the central part of Roraima State, often in sympatry with S. oyapockense s.l.
It has a very discrete distribution compared to the more widespread distribution of S.
oyapockense s.l. in this area. It is probable that some of the human-biting populations
recorded under S. oyapockense s.l. under “Material Examined” do pertain to S. roraimense
s.s because the two species are indistinguishable in the female. Taking into account the
two forms of this species from Ecuador and Argentina and the restricted distribution of S.
roraimense s.s., it is possible that the forms are cytospecies within a species complex.
Other species of the S. amazonicum group
Simulium chaquense and S. cuneatum are restricted to the north of Argentina and the
latter species has also been recorded in Paraguay; both species probably occur in Paraguay
and neighbouring Uruguay and southern Brazil. Similarly, S. ganalesense s.l. is restricted
to southern Mexico and Belize and S. sanguineum to the lowland forests of western
Colombia and Panama (Map 3). Simulium sanguineum is recorded as a vector of
Mansonella ozzardi in Colombia (Tidwell et al. 1980). Simulium venezuelense is
uncommon and only recorded from the northern Brazilian border area with French Guiana
and northern Venezuela (Map 3).
SIMULIUM AMAZONICUM
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MAP 3. The distribution of S. chaquense, S. cuneatum, S. ganalesense s.l., S. sanguineum, and S.
venezuelense of the amazonicum group in the Neotropical Region.
Discussion
From the preceding results we are drawing various conclusions based on the intraspecific
variation in various morphological characters. These are discussed under two sections:
pupae alone and link-reared adults.
PUPAE
The gill still remains the most reliable character for separation of pupae for species of
the amazonicum group. The number of filaments in the gill is consistent in almost all cases
and provides the first step in species identification. In only one locality cited in this paper
has gill number not resolved species identity where variation in female scutal patterns of
sympatric S. amazonicum s.l. and S. oyapockense s.l. was recorded. Gill length must be
used with caution because of the large variation in certain species and because of the
overlap of the variation in gill lengths between some of the different species. This is
particularly important in the two vector species of O. volvulus, S. oyapockense s.l. and S.
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SHELLEY ET AL.
roraimense s.s. Shelley et al. (1997) found a range of gill lengths of 0.7–1.5mm for S.
oyapockense s.l. and 1.9–2.6mm for S. roraimense s.s., which could be used to separate the
two species. However, they sounded a cautionary note that with higher sample size a
single range of lengths might occur. This has now been confirmed by our current data,
where S. oyapockense from the type locality has a gill length range of 1.6–2.1mm and S.
oyapockense s.l. from other localities (including the Bem Querer form on which the
description in Shelley et al. 1997 was based) of 0.8–1.9mm. Simulium roraimense
topotypes varied from 1.6–2.6mm and specimens from other localities from 1.4–2.0mm
(Table 1). Hence, length of gill cannot be used alone for separation of these two species.
Configuration of filament branching may be used for separation of several species, but
caution must be exercised because of variation even within the same specimen, well
illustrated in S. amazonicum (Figs. 17, 18).
TABLE 2. Synopsis of morphological characters of pupae of S. amazonicum group species.
Species
Gill
Gill length
Cephalothorax
Frontoclypeus
fila- Mean (range)
ments
Tubercles Trichome Tubercles
Trichome
branches
branches
venezuelense
4
-
(0.9–1.4) Mr, Mpor
argentiscutum
6
-
(0.8–0.9)
ganalesense (t.l.)
6
1.8 (1.6–2.1)
ganalesense
Belize
6
-
oyapockense s.l.
(t.l.)
6
oyapockense s.l.
(other)
Facial
Frontal
Spine
combs on
tergites 2,3
1,2,4
none
2,3
1
-,-
AMPr
1,2
r
1,2
1,2
-,-
(2.4-3.9)
AMpor
1,2
r
2
1,2
-,-
1.8
(1.6-2.1)
AMpor
1,2,3
r
1,2
1 (r)
+,+
6
-
(0.8-1.9)
AMpor
1,2,3
r
1,2
1 (r)
+,+ ;-,+ ;-,-
roraimense (t.l.)
6
1.8 (1.6-2.6)
AMPpor
1,2
por
1,2
1
-,-
roraimense (other)
6
-
(1.4-2.0)
AMPpor
1,2,3
r, por
1,2
1
-,-
amazonicum
8
1.4 (1.0-1.8)
AMPpor
1,2,3,4
por
2,3
1,2,3
-,-
chaquense
8
-
AMPpor
2
por
1
1
-,-
sanguineum
8
AMPpor
1,2,3
r
1,2
1
-,-
cuneatum
10
? por
1
Por
1
(1.8-2.0)
1.6 (1.5-1.7)
- (1.2-1.8)
1
?
Symbols as in Table 1
t.l. = type locality
The distribution and form of tubercles on the cephalothorax and frontoclypeus appear
to be species specific in most cases, but given the variation in density of tubercles on these
structures, we are reticent to use this character alone for identification (Table 2). There was
SIMULIUM AMAZONICUM
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no apparent sex difference in tubercle form and distribution. Tubercles were recorded on
all parts of the cephalothorax of S. amazonicum s.l., S. argentiscutum, S. roraimense s.l.,
and S. sanguineum, but only on the anterior and median areas in S. ganalesense and S.
oyapockense s.l. and median part of S. cuasisanguineum (synonym of S. oyapockense s.l.)
and S. venezuelense. Py-Daniel and Pessoa (2005) recorded a uniform distribution of
tubercles in species of their genera Cerqueirellum, Coscaroniellum, and Psaroniocompsa
[= subgenus Psaroniocompsa in this paper], but restricted to the posterior part of the
thorax in their new genus Shelleyellum [= siolii group of subgenus Psaroniocompsa in this
paper]. However, they noted the presence of small numbers of “small triangular
tubercles”, with no indication of thoracic distribution, in type material of Cerqueirellum
venezuelense [= amazonicum group of the subgenus Psaroniocompsa] that were recorded
as absent in the original description by Ramírez Pérez & Peterson (1981). Tubercles are
pointed and rounded in all species except S. argentiscutum, which only has rounded
tubercles. However, in populations of S. venezuelense, individuals were recorded in which
tubercles were either just rounded or both pointed and rounded (cf. Py-Daniel & Pessoa
2005). It is probable that with larger sample numbers similar variation would be observed
in other species and so this character would be of limited use in identification. Similarly,
the presence or absence of tubercles on the frontoclypeus is treated with caution as an
interspecific character. Absence of tubercles was recorded only in S. venezuelense (Table
2) in low sample numbers, but the validity of this character is still debatable. Other good
interspecific characters distinguish this species. This is not the case for S.
cuasisanguineum (a synonym of S. oyapockense s.l.), which is also devoid of tubercles.
This character, together with male scutal pattern and adult leg colour, was used by Ramírez
Pérez et al. (1982) for erecting S. cuasisanguineum as a new species. Shelley et al. (1987)
discounted the characters observed in the adults as interspecific and were in doubt about
the validity of the lack of tubercles in the pupa being an interspecific character because
insufficient material had been examined to properly assess variation in this character.
Consequently, they synonymised S. cuasisanguineum with S. oyapockense s.l. as an
interim measure until further material could be examined. We maintain this synonymy
because no other reliable characters are available for definition of this species and no
further specimens have been examined. Tubercle form in the frontoclypeus appears to be
generally consistent in most species with only rounded tubercles present in all species
except S. amazonicum s.l., S. chaquense, S. cuneatum, and S. roraimense s.s. where both
rounded and pointed tubercles are present. Shelley et al. (1997) discussed the use of
tubercle form for distinguishing between S. roraimense at its type locality and S.
oyapockense s.l. from a location some 100 km away, where only the former possessed
pointed tubercles on the frontoclypeus. Our present data show that only rounded tubercles
are present in S. oyapockense at its type locality and 16 other localities in northern Brazil,
but at the R. Jacy Paraná and Ituxi, both pointed and rounded are present. In S. roraimense
s.l., both pointed and rounded tubercles are present in all localities, except in one male
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SHELLEY ET AL.
from R. Catrimani and all specimens of the Manabi form from the R. Portoviejo, Ecuador
(Table 1), where the male scutal pattern is of the S. roraimense type (Figs. 137–140) and
the gill length falls within the ranges for both S. oyapockense s.l. and S. roraimense.
Hence, tubercle form may only be used for separation of S. oyapockense s.l. from S.
roraimense with caution, positive identification still relying on male scutal pattern.
Although reduced trichome length was seen only in populations of S. oyapockense s.l.,
this character failed to work in populations from the rivers Ajarani 1, Ituxi, Jacy Parana,
Madeira, and Toototobi (Table 1), where male scutal pattern indicated S. oyapockense s.l.
Again, this character may only be used with caution and in combination with other more
reliable characters.
Spine comb presence or absence on tergites II and III is also a weak character for two
reasons. Firstly, spine combs are very difficult to see, even in good preparations, and
secondly they were not always present on both segments in specimens from different
populations (Table 1).
In conclusion, based only on pupal characters, the gill filament number and in some
cases the branching configuration are reliable for species identification. Using a
combination of characters species identification can be more reliable, but given the
variations observed in small samples already discussed above, we recommend the rearing
of pupae to adults in most cases to be necessary before species identification is confirmed.
Additionally, several other species of simuliids that could occur sympatrically with species
of the amazonicum group have similar pupal gill form.
ADULTS
Based on the scutal pattern descriptions of females and males linked to pupal gill form
under “Results”, it is possible to recognise species of the amazonicum group that are listed
before. Some of these species show considerable morphological variation, which can be
interpreted only by recourse to cytological and molecular methods. Simulium oyapockense
s.l. and S. roraimense s.s. from Brazil have been studied in depth and shown to vary
intraspecifically in both pupal gill configuration and female scutal pattern. The variations
in pupal gill form (Figs. 13, 25, 26) and in female scutal pattern (Figs. 71–84) of S.
oyapockense s.l. cannot be linked to one another. The meaning of this variation in relation
to species limits requires further investigation. A similar situation is recorded here for S.
amazonicum s.l. along a short stretch of the R. Purus where two different female scutal
patterns occur, together with the typical pattern. Females of all three scutal pattern types
bite humans. The two types from the R. Ituxi (Figs. 67, 68) and R. Mari (Figs. 69, 70)
show no consistent difference in their pupal gill form to the basic pattern of S. amazonicum
s.s. (Fig. 8) and variations seen within topotypic material and two specimens from the R.
Ituxi (Figs.17–22). Since we are unable to interpret these variations as being intraspecific
or interspecific at this stage, we refer to both S. amazonicum and S. oyapockense as sensu
lato. Male scutal pattern also varies in S. roraimense. Males from the R. Portoviejo
SIMULIUM AMAZONICUM
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(Manabi form) in Ecuador have roughly square-shaped black cunae in the intervittal bands
(Fig. 139). Coscarón & Coscarón-Arias (2000) provided a description of S. oyapockense
s.l. from specimens collected in Salta, Tucumán, and Santiago del Estero provinces in
Argentina and the State of Amazonas, Brazil. The gill length of the specimens from
Argentina falls within the new ranges for S. oyapockense s.l. and S. roraimense s.l., but the
presence of long trichomes and the male scutal pattern indicates the species to be S.
roraimense s.l. We group S. roraimense s.s. together with these two forms under S.
roraimense s.l.
Distinction between the two human onchocerciasis vectors S. oyapockense s.l. and S.
roraimense s.l. is impossible in the female and with the newly found overlap in pupal gill
length, this previously used character is ineffective except in specimens with the shortest
gills in the range (below 1.4mm being S. oyapockense s.l.) and with the longest gills in the
range (above 2.1mm being S. roraimense s.l.). It is only in the male that separation of
species is possible in most cases by the form of the scutal pattern. In assigning names to
biting females, we have taken into account the distribution of link-reared material. This
has indicated that S. roraimense s.s. in Brazil has a limited and discrete distribution
compared to the more widespread distribution of S. oyapockense s.l. Hence, we have
placed all the populations referred to by Shelley et al. (1997) as S. oyapockense/
roraimense under S. oyapockense s.l. In countries bordering Brazil we consider humanbiting species in the BMNH collection as S. oyapockense s.l. because evidence for any
other identification is not available. A problem does arise in biting populations in southern
Brazil, Paraguay, and Uruguay because of their relative proximity to localities in
Argentina where anthropophilic populations of S. chaquense, S. cuneatum, and S.
roraimense Argentina form [as S. oyapockense s.l. in Coscarón & Coscarón-Arias 2000]
occur. It is possible that females of one or more of these species could have been grouped
under S. oyapockense s.l. A further complication arises in that S. quadristrigatum, only
known from the female, is only recorded from this area. Simulium quadristrigatum was
described from “Parana” by Enderlein (1934) from a presumed human-biting, holotype
female (Figs. 59, 60) (see Hernández & Shelley 2005). These authors followed Vulcano
(1967) in supposing that this referred to Paraná State, Brazil, and not the view of Coscarón
(1983) that it could refer to the R. Paraná in either Brazil or Argentina. In this paper, we
are now following Coscarón’s view that “Paraná” could refer to either country and add
Paraguay to this list. These authors also agreed with Coscarón’s (1983, 1987) treatment as
a species inquirenda. However, as a result of this current study, we maintain S.
quadristrigatum as a valid species that cannot be fully defined because its type locality is
imprecise and topotypes cannot be collected. Based on its scutal pattern and provenance it
is closely related to S. chaquense, S. cuneatum, S. roraimense Argentina form, and S.
oyapockense s.l. from Brazil [see Maps 1–3]. Since the pupa of S. quadristrigatum is not
known, molecular examination of these species will be necessary in order to establish
which, if any, is a junior synonym of S. quadristrigatum, which predates all the other
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SHELLEY ET AL.
names. The consequence of a new synonymy in the case of S. chaquense or S. cuneatum is
straightforward. However, should S. oyapockense fall as a synonym to S. quadristrigatum,
serious complications would result in relation to the numerous publications concerning S.
oyapockense as a vector of O. volvulus. A new name would need to be provided for the
vector species. This will probably be the case anyway because we suspect the presence of
a species complex in S. oyapockense consisting of the zoophilic S. oyapockense s.s. from
the R. Oyapock and at least one other cytospecies consisting of the anthropophilic
populations found at Cachoeira Bem Querer and localities in the adjacent onchocerciasis
focus in Brazil. Further studies on link-reared specimens are necessary, integrating
morphological, cytological, and molecular parameters before the situation is resolved.
Preliminary cytological evidence for the possible existence of a species complex in S.
oyapockense s.l. was provided by Procunier et al. (1987), who distinguished the Manabi
form in Ecuador, collected from the R. Portoviejo, from S. roraimense s.s. and S. limbatum
in Brazil. This was based on differences in sex chromosomes and the centromere region in
chromosome I, but no banding sequences in the polytene chromosome maps were given.
In a subsequent paper, Procunier (1989) linked S. oyapockense Manabi form with S.
limbatum, S. quadrifidum, and S. sanguineum by two chromosomal centromere
dimorphisms in chromosomes I and II. He recorded the presence of a complex for S.
oyapockense consisting of the Manabi form from Ecuador and the Bem Querer form from
Brazil and “subsumed” S. roraimense within S. oyapockense s.l. Paradoxically, he
considered S. limbatum to be more closely related to S. roraimense than to S. oyapockense,
whereas based on morphological characters the latter two species are more closely related.
With our latest data analysis, we are placing the “Manabi form” of S. oyapockense s.l. with
S. roraimense s.s. under S. roraimense s.l. and include the Argentina form of S. roraimense
(Coscarón & Coscarón-Arias 2000) until cytological or molecular evidence indicates
whether these forms are different species from S. roraimense in Brazil.
Sympatry of three members of the amazonicum group (Maps 1, 2; Material Examined)
has produced a situation that cannot be resolved at this juncture. In the case of S.
oyapockense s.l. and S. roraimense s.s., the species appear to be good because distinct
scutal patterns occur in males collected from the same locality; no intermediate patterns
have been seen. The same does not apply to S. amazonicum s.l. where it is sympatric with
S. oyapockense s.l. at the R. Ituxi, a tributary of the R. Purus. In this area south of the R.
Amazon, S. amazonicum s.l. is the predominant species and S. oyapockense s.l.
(predominant north of the R. Amazon) is far less common. We have recorded a female
identified as S. amazonicum s.l. based on an 8-filamented pupal gill with a scutal pattern
(Figs. 67, 68) resembling that of S. amazonicum s.s. (Figs. 47, 48) yet with intervittal
cunae similar to S. oyapockense s.s. (Figs. 57, 58). This was in sympatry with a typical S.
oyapockense s.l. reared from a 6-filamented pupa and a typical female scutal pattern (Figs.
79, 80), and a specimen with a typical male scutal pattern was reared from a 6-filamented
pupa. Further study is required in this area to clarify the situation.
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Nomenclatural rearrangements involving the amazonicum and siolii groups
1274
We include comments on the siolii group here because we have dealt with S. venezuelense,
included in the amazonicum group by Crosskey & Howard (2004), and discuss (below) S.
damascenoi of the siolii group as its possible synonym. The siolii group contains species
that appear to fit in the auristriatum group (S. guaporense) and the amazonicum group (S.
tergospinosum and S. siolii) if scutal pattern is considered as the main species group
character. However, the form of the male gonostyle justifies the continuing existence of
the siolii group. A similar situation occurs in the subgenus Psilopelmia where scuta are
typically orange, yet S. haematopotum and S. quadrivittatum that pertain to this subgenus
resemble amazonicum group species in their scutal patterns. The diagnostic characters of
the siolii group are that the larvae have fan-like setae and dorsal protuberances (Py-Daniel
1989). There is some controversy over the location of these dorsal protuberances (Hamada
et al. 2006). We have comments on the validity of two of the species and the assignment of
species to the siolii group. In the case of S. damascenoi, Py-Daniel (1988) provided
descriptions and figures of the female morphology from two specimens removed from
pharate pupae, and of the pupa and the larva collected in Amapá state. He also provided a
key to larvae of species of Psaroniocompsa whose pupae have gills with four filaments (S.
damascenoi, S. lourencoi, S. siolii, and S. schmidtmummi) and a key to the pupae of these
species. In his 1989 paper, Py-Daniel provided tables to distinguish larvae and pupae of
the siolii group, including the three former species and S. guaporense. In this 1989 paper,
he referred in Table 2 to rounded tubercles on the frontoclypeus of S. damascenoi, which
were described correctly as being absent in his previous paper (Py-Daniel 1988). No
mention was made of S. venezuelense, which has a pupal gill with four filaments and was
included in the amazonicum species group of the subgenus Psaroniocompsa (see Shelley
1988) prior to its transfer to the previously erected subgenus Cerqueirellum (Py-Daniel
1983). In 2000, Hamada described a new species, S. tergospinosum, from southern
Amazonas state in Brazil, which she placed in the siolii group at the same time confirming
Py-Daniel’s (1989) diagnosis of the species group. She also compared S. tergospinosum
with S. damascenoi, but in the case of the latter species not with other members of the
siolii group. In 2003, Hamada et al. described the male of S. damascenoi as well as
additional female characters from material collected at its type locality and adjacent areas
due to the non-availability of type material. We have examined link-reared specimens
from this area of Brazil and compared these with the descriptions of female, pupa, and
larva of S. damascenoi of the previously cited authors as well as the female holotype, a
reared female and exuviae, and three larval paratypes and regard our specimens as
conspecific (see Material Examined). However, the gonostyle of the male recorded in this
current paper as S. venezuelense differs from that described and figured as S. damascenoi
by Hamada et al. (2003). Their figure shows a longitudinal ridge shared by other species
of the siolii group, but the gonostyle appears shorter and reminiscent of some species of
the amazonicum group. The gonostyle on our specimens is typical in form for the
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SHELLEY ET AL.
amazonicum group, but has a very slightly defined ridge. We also compared our material
with descriptions by the original authors of S. venezuelense (for which no larval
description was made) because no type material was available for examination. We accept
our material collected in the same area of Brazil from which S. damascenoi was described
as being conspecific with S. venezuelense from Venezuela because no morphological
differences were observed in major interspecific characters including the male gonostyle.
We are hesitant to synonymise S. damascenoi with S. venezuelense because of the apparent
differences in gonostyle morphology. It is not yet clear whether the difference in gonostyle
morphology between the drawn figures of Hamada et al. (2003) and our digitally imaged
figure is due to different specimen orientation and recording methods or is a true
difference. In 1988, Py-Daniel described S. lourencoi from larvae and pupae collected in
southern Amazonia, Brazil. He distinguished the larva of this species from that of S. siolii
based on the number of rays on the cephalic fan, different antennal segment proportions,
number of hooks per row on the posterior sucker, the intensity of the positive head pattern
and difference in number of lobules on the anal lobes. We have examined the pupal
holotype and three pupal paratypes. We consider these characters to be very variable
within populations and dependent on specimen size. In the pupae, the configuration of gill
branching and the presence or absence of sclerotised abdominal striations were used to
distinguish the species, but we consider these differences as intraspecifically variable.
Consequently, we are synonymising S. lourencoi with S. siolii. We consider the use of
larval tubercles and morphology of larval setae as characters for forming species groups of
insufficient weight. The former vary from absent to well developed even at the level of
cytospecies. Similar variation also occurs in the S. damnosum species complex in Africa
and in the S. guianense species complex from Brazil (Shelley et al. 2002). The recent
elevation of the siolii group to the genus Shelleyellum by Py-Daniel & Pessoa (2005), of
which the type species is S. siolii (Py-Daniel & Pessoa 2006), is not accepted because the
characters that they used to separate the siolii group from other species groups in the
subgenus Psaroniocompsa are of insufficient weight to justify generic status.
Consequently, Shelleyellum becomes a junior synonym of Simulium Latreille and the siolii
group remains within the subgenus Psaroniocompsa of the genus Simulium. Hamada et al.
(2006) regarded S. guaporense as unassigned to species group within the subgenus
Psaroniocompsa because its scutal pattern does not resemble other species in the group.
Acknowledgements
The first author would like to thank the Leverhulme Trust for an Emeritus Fellowship that
allowed the completion of this work. We also acknowledge funding over the years from
the Natural History Museum, London, and the Oswaldo Cruz Institute, Brazil, for
collecting and curating Neotropical simuliids. We thank Dr R.W.Crosskey for his advice
on nomenclatural problems and Anne Shelley for her organization of the figures and
SIMULIUM AMAZONICUM
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production of the maps. Luis M. Hernández would like to thank Victor Py-Daniel and
Neusa Hamada from the Instituto Nacional de Pequisas da Amazônia (INPA) for the loan
of material housed in INPA collections. We also acknowledge an anonymous referee for
valuable comments on the relationship between S. damascenoi and S. venezuelense.
References
Coscarón, S. (1971) Notas sobre simulidos Neotropicales 1. Sobre una nueva especie del norte
argentino (Diptera, Simuliidae). Revista de la Sociedad Entomológica Argentina, 33, 33–41.
Coscarón, S. (1983) Notas sobre simulidos Neotropicales XII. Las especies de Simulium “grupo
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Sociedad Entomológica Argentina, 89–99.
Coscarón, S. (1987) El género Simulium Latreille en la Región Neotropical: Análisis de los Grupos
Supraspecíficos, Especies que los Integran y Distribución Geográfica (Simuliidae, Diptera).
Museu Paraense Emílio Goeldi, 112 pp.
Coscarón, S. (1991) Fauna de agua dulce de la Republica Argentina. 38. Insecta Diptera Simuliidae. Fasciculo 2. [vi] 7–304 + 78 unnumbered pages of figures and legend. Fundación para la
Educación, La Ciencia y la Cultura, Buenos Aires, Argentina.
Coscarón, S. & Coscarón-Arias, C.L. (2000) New species records for the blackfly (Diptera-Simuliidae) fauna of Argentina with description of adults, pupa and larva of Simulium oyapockense
s.l. and S. seriatum. Memórias do Instituto Oswaldo Cruz, 95, 179–187.
Crosskey, R.W. & Howard, T.M. (2004) A Revised Taxonomic and Geographical Inventory of
World Blackflies (Diptera: Simuliidae). The Natural History Museum, London, 82 pp. [web
version: http://www.nhm.ac.uk/research-curation/projects/blackflies/Intro.pdf]
Enderlein, G. (1934) Weiterer Ausbau des systems der Simuliiden (Dipt.). Deutsche Entomologische Zeitschrift, 1933, 273–292.
Enderlein, G. (1936) Simuliologica 1. Sitzungsberichte der Gesellschaft Naturforschender Freunde
Berlin, 1936, 113–130.
Floch, H. & Abonnenc, E. (1946) Simulidés de la Guyane Française (II). S. cauchense n.sp., S.
oyapockense n.sp., S. iracouboense n.sp. Publication de l’Institut Pasteur de la Guyane et du
Territoire de l’Inini, 137, 1–19.
Goeldi, E.A. (1905) Os mosquitos no Pará. Memórias do Museu Goeldi (Museu Paraense) de Historia Natural e Ethnographia, 4, 138–139.
Hamada, N. (2000) Simulium (Psaroniocompsa) tergospinosum new species (Diptera: Simuliidae)
in siolii group from the southern part of the state of Amazonas, Brazil. Memórias do Instituto
Oswaldo Cruz, 95, 819–828.
Hamada, N. & Fouque, F. (2001) Black flies (Diptera: Simuliidae) of French Guiana: Cytotaxonomy and a preliminary list of species. Memórias do Instituto Oswaldo Cruz, 96, 955–959.
Hamada, N., Ale-Rocha, R. & Luz, S.L.B. (2003) description of Simulium damascenoi (Diptera:
Simuliidae) male and the black-fly species from the state of Amapá, Brazil. Memórias do Instituto Oswaldo Cruz, 98, 353–360.
Hamada, N., Hernández, L.M. & Luz, S.L.B. (2006) Taxonomy of Simulium guaporense Py-Daniel
(Diptera: Simuliidae) from Brazil, with the first description of males and females. Zootaxa,
1104, 23–34.
Nunes de Mello, J. A. S. (1974) Simulídeos (Diptera, Nematocera) do Território Federal de
Roraima (Brasil). Doctoral thesis, Faculdade de Medicina, Universidade de Sorocaba, São
Paulo, Brasil. 56 pp.
Procunier, W.S. (1989) Cytological approaches to simuliid biosystematics in relation to the epide46
SHELLEY ET AL.
miology and control of human onchocerciasis. Genome, 32, 559–569.
Procunier, W.S., Shelley, A.J. & Arzube, M. (1987) Cytological identification of Simulium
oyapockense Manabi form (Diptera: Simuliidae): a potential vector of onchocerciasis in Ecuador. Tropical Medicine and Parasitology, 38, 71.
Py-Daniel, V. (1983) Caracterização de dois novos subgéneros em Simuliidae Diptera: Culicomorpha) Neotropical. Amazoniana, 8, 159–223.
Py-Daniel, V. (1988) Simuliidae (Diptera: Culicomorpha) no Brasil. VI. Sobre Simulium (Psaroniocompsa) siolii sp.n., Simulium (P.) lourencoi sp.n., e Simulium (P.) damascenoi sp.n. Revista de
Saúde Pública, 22, 292–310.
Py-Daniel, V. (1989) Simuliidae (Diptera: Culicomorpha) no Brasil. X – Descrição de Simulium
(Psaroniocompsa) guaporense sp.n. Revista de Saúde Pública, 23, 502–508.
Py-Daniel, V. & Pessoa, F.A.C. (2005) Simuliidae (Diptera: Culicomorpha) no Brasil XI – Shelleyellum gênero novo. Acta Amazonica, 35, 473–486.
Py-Daniel, V. & Pessoa, F.A.C. (2006) Designação da espécie-tipo de Shelleyellum Py-Daniel &
Pessoa, 2005 (Diptera: Simuliidae). Acta Amazonica, 36,133.
Py-Daniel, V. & Moreira Sampaio, R.T. (1994) Jalacingomyia gen.n. (Culicomorpha); ressurreição
de Gymnopaidinae; a eliminação do nível tribal; apresentação de novos caractéres e a
redescrição dos estágios larval e pupal de Simulium colombaschense (Fabricius, 1787)
(Diptera: Simuliidae). Memórias del CAICET, 4, 101–148.
Py-Daniel & Moreira Sampaio, R.T. (1995) Gêneros e espécies de Simuliidae (Diptera: Culicomorpha) assinalados para o Brasil até 1995. Entomologia y Vectores, 2, 117–121.
Ramírez Pérez, J. (1971) Distribución geográfica y revisión taxonómica de los simúlidos (Diptera:
Nematocera) de Venezuela con descripción de diez especies nuevas. Acta Biológica Venezolana, 7, 271–371.
Ramírez Pérez, J. (1983) “Los jejenes de Venezuela.” Simposio de oncocercosis Americana. Caicet,
Puerto Ayacucho, 15–17 Octubre, 1983, iii + 156 pp.
Ramírez Pérez, J. & Peterson, B. (1981) Estudio del complejo Simulium amazonicum-sanguineum
en Venezuela. Descripción de tres nuevas especies. Boletin de la Dirección de la Malariologia
y Saneamiento Ambiental, 31, 151–160.
Ramírez Pérez, J., Yarzábal, L. & Peterson, B. (1982) La simuliofauna del Territorio Federal Amazonas (Venezuela). Publicación Científica 1, 1–104, Proicet, Amazonas.
Shelley, A.J. (1988) Biosystematics and medical importance of the Simulium amazonicum group
and the S. exiguum complex in Latin America. In: Service, M.W. (Ed.), Biosystematics of Haematophagous Insects, Oxford University Press, Oxford, pp. 203–220.
Shelley, A.J. (2002) Human onchocerciasis in Brazil: an overview. Cadernos de Saúde Pública, 18,
1167–1177.
Shelley, A.J. & Coscarón, S. (2001) Simuliid blackflies (Diptera: Simuliidae and ceratopogonid
midges (Diptera: Ceratopogonidae) as vectors of Mansonella ozzardi (Nematoda: Onchocercidae) in northern Argentina. Memórias do Instituto Oswaldo Cruz, 96, 451–458.
Shelley, A.J., Hernández, L.M., & Davies, J.B. (2004) Blackflies (Diptera: Simuliidae) of southern
Guyana with keys for the identification of adults and pupae – a review. Memórias do Instituto
Oswaldo Cruz, 99, 443–470.
Shelley, A.J., Hernández, L.M. & Penn, M. (2002) A biosystematic revision of the blackflies
(Diptera:Simuliidae) of Belize, Central America. Bulletin of the British Museum (Natural History) Entomology series, 71, 135–271.
Shelley, A.J. & Luna Dias, A.P.A (1980) Simulium argentiscutum sp. nov. Diptera: Simuliidae), a
member of the S. amazonicum—group of species: description of adults, pupa and larva.
Memórias do Instituto Oswaldo Cruz, 75, 105–111.
Shelley, A.J., Lowry, C.A., Maia-Herzog, M., Luna Dias, A.P.A. & Moraes, M.A.P. (1997) Biosystematic studies on the Simuliidae (Diptera) of the Amazonia onchocerciasis focus of Brazil.
SIMULIUM AMAZONICUM
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Bulletin of the British Museum (Natural History) Entomology series 66, 1–121.
Shelley, A.J., Luna Dias, A.P.A., Maia-Herzog, M., Procunier, W.S. & Moraes, M.A.P. (1987) Identification of vector species (Diptera: Simuliidae) of human onchocerciasis in the Amazonia
focus of Brazil and Venezuela. Memórias do Instituto Oswaldo Cruz, 82, 461–465.
Shelley, A.J., Pinger, R.R. & Moraes, M.A.P. (1982) The taxonomy, biology and medical importance of Simulium amazonicum Goeldi (Diptera: Simuliidae), with a review of related species.
Bulletin of the British Museum (Natural History) Entomology series, 44, 1–29.
Tidwell, M.A., Peterson, B.V., Ramírez Pérez, J., de Tidwell, M., Lacey, L.A. (1980) Notas y claves
preliminaries de los jejenes neotropicales pertenecientes a los grupos Simulium amazonicum y
S. sanguineum (Diptera: Simuliidae) incluyendo los vectores de Onchocerca volvulus y Mansonella ozzardi. Boletin de la Direccion de Malariologia y Saneamiento Ambiental, 21, 79–89.
Tidwell, M.A., Tidwell, M.A. & Peterson, B.V. (1981a) A redescription of the female of Simulium
sanguineum Knab and descriptions of the male, pupa and larva (Diptera: Simuliidae). Proceedings of the Entomological Society of Washington, 83, 13–27.
Tidwell, M.A., Peterson, B.V., Ramírez Pérez, J., Tidwell, M. & Lacey, L.A. (1981b) Notas y
claves preliminaries de los jejenes Neotropicales pertenecientes a los grupos Simulium amazonicum y S. sanguineum (Diptera:Simuliidae) incluyendo los vectores de Onchocerca volvulus y Mansonella ozzardi. Boletin de la Direccion de Malariologia y Saneamiento Ambiental,
21, 79–89.
Vargas, Martínez Palácios & Díaz Nájera, A. (1946) Simúlidos de México. Revista del Instituto de
Salubridad y Enfermedades Tropicales, 7, ii + 101–192 + 25 plates of figures on un-numbered
pages and legends to these figures, also on un-numbered pages.
Vulcano, M.A. (1967) A Catalogue of the Diptera of the Americas South of the United States. 16.
Family Simuliidae. Departamento de Zoologia, Secretaria de Agricultura, Sno Paulo, Brazil. 44
pp.
World Health Organization (1979) Report of an informal workshop on the taxonomy of South
American Simuliidae of medical importance. Fundação Oswaldo Cruz, Rio de Janeiro, Brazil,
7–11 May 1979. Mimeographed document. TDR/FIL/SIM/79.1. 39 pp.
World Health Organization (1982) Report of an informal workshop on the taxonomy of South
American Simuliidae of medical importance. Fundação Oswaldo Cruz, Rio de Janeiro, Brazil,
16–26 November 1982. Mimeographed document. TDR/FIL/SIM/82.3. 13 pp.
48
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Appendix
ZOOTAXA
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Material examined
Abbreviations of depositories
AMNH
BMNH
INPA
IOC
NMHU
SMT
American Museum of Natural History, New York, USA.
The Natural History Museum, London, UK.
Instituto Nacional de Pesquisas da Amazônia, Brazil
Instituto Oswaldo Cruz, Rio de Janeiro, Brazil.
Museum für Naturkunde der Humboldt-Universität, Berlin, Germany. [Previously cited
as Zoologisches Museum der Humboldt-Universität, Berlin, Germany in Shelley et al.,
1997.].
Staatliches Museum für Tierkunde, Dresden, Germany.
Simulium (Psaroniocompsa) amazonicum Goeldi
All material listed under Shelley et al. (1982) was examined for this paper. Specimens that have
been subsequently dissected, imaged, and to which new labels have been added, as well as further
material examined are as follows:
TYPE MATERIAL
BRAZIL
Amazonas State
PINNED
Valparaiso, R. Purus; 22.xi.1977, (A.J.Shelley)—1& (reared, pupa on slide) [NEOTYPE]
(BMNH, BM 1979-580) [The specimen bears several labels: 1- White, red-bordered round label “
NEOTYPE”; 2- White label with locality information; 3- White label with date, collectors name
and BMNH number); 4- White handwritten label “Valparaiso 2 km from site of Bom Lugar, see
Shelley, Pinger, Moraes 1982, Bull. Mus. Nat. Hist. Vol. 44(1), p.9; 4- White label “Simulium
amazonicum, det. A.J.Shelley; 6- White label “Digital Photograph BMNH (E)# 625002- & (reared,
pupa on slide) (BMNH).
SLIDE
Same label details as pinned neotype—pupal exuviae [Neotype] (BMNH).
OTHER MATERIAL
BRAZIL
Amazonas State
PINNED
R. Purus, Bom Lugar; [Without date.], (J. Huber)—1& (BMNH, pres. by E. A. Goeldi). [One
of original “cotypes” not recorded in Shelley et al. (1982) but subsequently found.] Same locality as
SIMULIUM AMAZONICUM
49
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1274
Neotype, (site 277), 29–30.xi.1976; (sites 293, 294), 17–19.vi.1977; (site 280), 3.xii.1977, (W.
Arouck)—numerous females (man-biting) (BMNH, B.M. 1979-580). Bom Lugar, R. Purus, (site
279); 2.xii.1976, (W. Arouck)—7&& (man-biting) (BMNH; B.M. 1979-580); (site 292), 19.vi.1977,
(W. Arouck)—5&& (man-biting); (site 323-12), 22.xi.1977, (A.J.Shelley)—1% (reared) (BMNH,
B.M. 1979-580). Valparaiso, R. Purus, (site 323-10); 22.xi.1979, (A.J.Shelley)—1% (reared)
(BMNH, BM 1979-580). Valparaiso, R. Purus, (site 323-5-7); 22.xi.1979, (A.J.Shelley)—2&&
(reared); 1& only thorax (head, wings, legs, abdomen, and pupal skin on slide); 1& only thorax
(head, legs, one wing and genitalia on slide). Lábrea, Lábrea-Humaita Rd (Transamazônica), 30 km
after Lábrea, R. Mari, (site 1411); 9.viii.1999, (A.J.Shelley & A. P. Luna Dias)—20&& (man-biting)
(BMNH; IOC). Lábrea, Palmeri (Minaã), R. Ituxi, (site 1412), 07º19’S64º56’W; 9.vii.1999,
(A.J.Shelley & A.P.A. Luna Dias)—11&& 10%% (reared) (BMNH; IOC); 1& (man-biting) (BMNH).
R. Ituxi, Igarapé do Paraná, (site 269); 13—30.ix.1976, (A.J.Shelley)—3&& (reared, pupa on slide),
5&& (man-biting), 1% (reared) (BMNH). Pauini, Fazenda Pauini, R. Pauini, (site 1409); 4.viii.1999,
(A.J Shelley & A.P.A. Luna Dias)—16 && (man-biting), 3%% (reared) (BMNH; IOC).
SLIDE
Bom Lugar, R. Purus, (No. 292), 19.vi.77, (No. 276), 29.ii.77, (W. Arouck)—several slides with
&, %, pupae and larvae, the & collected from human bait and horse (BMNH). Valparaiso, R. Purus,
(site 323-5-7); 22.xi.1979, (A.J.Shelley)—2&& (reared) 1& head, wings, legs, abdomen, and pupa
(thorax pinned); 1& head, legs, one wing; and genitalia (thorax pinned). Bom Lugar, R. Purus, (site
323); 22.xi.1977, (A.J.Shelley)—1& (man-biting) (BMNH).
SPIRIT
Bom Lugar, (site 293, 295), 8º42’S 67º22’W; 18.vi.1977, (W. Arouck)—several vials with
numerous && (man-biting); (site 277), 30.xi.1976—1 vial with numerous && (man-biting)
(BMNH); (site 294), 17.vi.1977, (W. Arouck)—3 vials with numerous && (man-biting). Bom
Lugar, (site 323), 22.xi.77, (A.J.Shelley)—2 vials with numerous && (man-biting) (BMNH); 17—
19.vi.1977 (W. Arouck)—2 vials with numerous && (man-biting) (BMNH). Capacini, R. Purus (site
86), 10.x.1973, (A.J.Shelley)—4 vials with numerous && (man-biting) (BMNH). Feijoal, R.
Solimões, (site 392); 5.x.1978, (A.J.Shelley & A.P.A.Luna Dias)—1 vial with 8&& (man-biting)
(BMNH, B.M.1979-580). R. Ituxi, Igarapé do Paraná, (site 269); 13—30.ix.1976, (A.J.Shelley)—4
vials with && and pupae (BMNH, B.M.1979-580). Lábrea, Lábrea-Humaita Rd (Transamazônica),
30 km after Lábrea, R. Mari, (site 1411); 8.viii.1999, (A.J.Shelley & A. P. A. Luna Dias)—
numerous && (man-biting) (BMNH).
Roraima State
SPIRIT
Roraima Territory, Near N. Perimeter Road, R. Repartimento, (site 447), 28.iv.1979,
(A.J.Shelley)—numerous && (man-biting) (BMNH, B.M. 1979-580). Roraima Territory, Near N.
Perimeter Road, R. Ajarani 1, (site 22); 16.i.1979, (A.J.Shelley & A.P.A.Luna Dias)—8&& (manbiting), 3 pupae (BMNH, B.M. 1979-580).
Simulium (Psaroniocompsa) argentiscutum Shelley & Luna Dias, 1980
50
SHELLEY ET AL.
All material listed under Shelley & Luna Dias (1980) was examined for this paper. Specimens that
have been subsequently dissected, imaged, and to which new labels have been added, as well as
further material examined are as follows:
TYPE MATERIAL
BRAZIL
Rondônia State
PINNED
Cachoeira Teotônio, R. Madeira, (site 398-5); 10.x.1978, (A.J.Shelley & A.P.A.Luna Dias) —1
& (reared) [HOLOTYPE] (IOC, B.M. 1979-580) [The specimen bear several labels: 1- White, redrounded labels “Holotype”; 2- White label with locality information; 3- White label with collector’s
name and BM number; 4- White label “S. argentiscutum”; 5- White label “S. argentiscutum Shelley
& Luna Dias Det. A.J.Shelley 1979 FioCruz No. 398-5” 6- White label “Digitally photographed
Th. ant. + post. + abdomen+ pupa by L.M.Hernández (BMNH 2004). [The thorax and the abdomen
of the holotype are in good condition, but the head and right wing are missing; only the right front
leg and hind leg are on the specimen, remaining legs missing.]. Same data as holotype—1%
(reared), only head and thorax, and pupal exuviae in glycerine (abdomen, legs, wings and genitalia
on slide). [PARATYPE] (BMNH). Same locality data as holotype—2&& (reared), only thorax and
pupa (head, wings, legs, abdomen and genitalia on slide) [PARATYPE] (BMNH, B.M. 1979-580).
SLIDE
Same data as holotype—1% (reared); only abdomen, legs, wings and genitalia (head and thorax
pinned, pupal exuviae in glycerine) [PARATYPE] (BMNH). Same locality date as holotype—1%
(reared; only head and thorax; abdomen, legs, wings and genitalia on slide) [PARATYPE]
(BMNH). Same locality data as holotype—2&& (reared), only head, wings, legs, abdomen and
genitalia (thorax pinned, pupa in glycerine) [PARATYPE] (BMNH, B.M. 1979-580).
As Simulium urubambanum Enderlein
PERU
PINNED
Unini, R. Ucayali; 20.x.1903, [Without collector’s name.]—1& [PARATYPE] (SMT) (only
thorax, abdomen and legs; head on slide) [The specimen bears several labels: 1- Green label with
locality date; 2- Orange label “TYPUS”; 3- White label with Enderlein handwriting “Simulium
urubambanum type Enderl. &”, printed “Dr. Enderlein det. 19[33]”; 4- White label “Paratype of
Simulium urubambanum Enderlein, 1924, examined by A.J.Shelley 1988”; 5- White label:
Simulium argentiscutum Shell. & L. D. det. A.J.Shelley.]
OTHER MATERIAL
BRAZIL
Amazonas State
PINNED
SIMULIUM AMAZONICUM
51
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Tefe, R.Solimões; 22.vii.1976, (A.J.Shelley)—5&& (man-biting) (BMNH, B.M. 1979-580).
Tauaru, Benjamin Constant, Rio Solimões; 20.viii.1961, (Eduardo)—2&& (BMNH). Ig. Codajás;
21.viii.1958, (Antonio)—1& (not associated with pupa) (BMNH).
Rondônia State
PINNED
Guajará Mirim; 5.ix.1963, 13.vi.1961, (Eduardo)—2&& (reared, but not associated with
pupae) (BMNH).
Simulium (Psaroniocompsa) chaquense Coscarón
TYPE MATERIAL
ARGENTINA
PINNED
Chaco Province, Nr. Formosa, Arroyo Zapirain; 14.vii.1971, (S. Coscarón)—1& (reared), only
thorax (head, wings, legs, abdomen, genitalia, and pupal exuviae on two slides) [PARATYPE]
(BMNH); 1% (reared, pupa on slide) [PARATYPE] (BMNH).
SLIDE
Chaco Province, Nr. Formosa, Arroyo Zapirain; 14.vii.1971, (S. Coscarón)—1& (reared), only
head, wings, legs, abdomen, genitalia, and pupal exuviae on two slides (thorax pinned); 1 pupal
exuviae (male pinned) [PARATYPE] (BMNH). Chaco Province, Nr. Formosa, Arroyo Carolis;
8.xi.1980, (S. Coscarón)—1& pupal pelt [PARATYPE] (BMNH).
OTHER MATERIAL
ARGENTINA
PINNED
Buenos Aires, Junin; xi or xii.1991, [Without collector’s name.]—1& (BMNH). Buenos Aires,
Junin, Brazo Norte; 23.xii.1991, (Marino)—1& (BMNH). Junin; 1991; (Marino)—1% (BMNH).
Corrientes, Bella Vista Est. Piscicultura; 14.x.1974, (Coscarón)—2&& (NMHU), 1& (BMNH) [All
specimens bear an identification label in Coscarón’s handwriting “Simulium chaquense det. S.
Coscarón 74”.]. Chaco, Ao. Carolis; 8.xi.1980, (Coscarón)—1& (reared) only thorax (head, one
wing, one leg, and pupa on slide); 1% (reared) only thorax and pupal exuviae; (wing, legs,
abdomen, and genitalia on slide; head, one wing, and two legs missing); 1% (reared); only head,
thorax, one fore leg, and pupal exuviae (wings, abdomen, remaining legs, and genitalia on slide).
Tres Rios, Concepción del Uruguay, Arroyo Itape; 5.xi.1980, (A.J.Shelley)—38&& (man-biting)
(BMNH).
SLIDE
Chaco, Ao. Carolis; 8.xi.1980, (Coscarón)—1& (reared); only head, one wing, one leg, and
pupa (rest of adult pinned); 1% (reared) only wing, legs, abdomen, and genitalia (thorax and pupal
52
SHELLEY ET AL.
exuviae pinned; head, one wing, and two legs missing); 1% (reared) only wings, abdomen, legs, and
genitalia (head, thorax, one fore leg, and pupal exuviae pinned).
SPIRIT
Buenos Aires, Rio Salado; 21.v.2003, (S. Coscarón)—several adults and pupae (BMNH).
Simulium (Psaroniocompsa) cuneatum (Enderlein, 1936)
TYPE MATERIAL
PARAGUAY
PINNED
[Without date or locality.]; (S.V. Fiebrig)—% (only head, thorax and left wing; rest on slide).
[HOLOTYPE] (NMHU), [The specimen bears several labels. 1- Green label, with country and
collector’s name; 2- Orange label, “Typus”; 3- White, hand-written label by Enderlein
“Psaroniocompsa cuneata Type [underlined in red] Enderl, % Dr. Enderlein det. 1934; 4- White,
red-bordered round label “Holotype”.]
SLIDE
Same data as pinned adult—1% (only right wing, five legs [1 missing], abdomen, and genitalia;
head, thorax, and left wing pinned). [HOLOTYPE] (NMHU).
OTHER MATERIAL
ARGENTINA
PINNED
Corrientes, R. Paraná, Costa Chaqueña; 28.x.1981, (S. Coscarón)—1& (reared) only thorax
and pupal exuviae (head, wings, legs, abdomen, genitalia on slide) (BMNH). Corrientes, R. Paraná,
Costa Chaqueña, frente a Valencia; 29.x.1981, (S. Coscarón)—1% (reared) only head, thorax, three
legs, and pupal exuviae (abdomen, genitalia, three legs, and wings on slide) (BMNH).
SLIDE
Corrientes, R. Paraná, Costa Chaqueña; 28.x.1981, (S. Coscarón)—1& (reared) only head,
wings, legs, abdomen, and genitalia (thorax and pupal exuviae pinned) (BMNH). Corrientes, R.
Paraná, Costa Chaqueña, frente a Valencia; 29.x.1981, (S. Coscarón)—1% (reared) only abdomen,
genitalia, three legs, and wings (head, thorax, three legs, and pupa) (BMNH).
SPIRIT
Corrientes, R. Paraná, Costa Chaqueña; 28.x.1981, (S. Coscarón)—several adults and pupae
(BMNH).
SIMULIUM AMAZONICUM
53
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Simulium (Psaroniocompsa) damascenoi Py-Daniel, 1988
TYPE MATERIAL
BRAZIL
Amapá State
SLIDE
Ig. do Enrique, Macapá-Oiapoque, km 290; 23.xii.1987, (Py-Daniel)—1& (reared)
[HOLOTYPE] (INPA) [The slide has several labels: On one side of the slide there is a white label
with locality, date and collector’s name. It also has handwritten on the label “INPA-6186-2 1 pupa 1
fêmea 6186”. On the other side it has a red, handwritten label “HOLOTIPO S. damascenoi &
(6186) and in black ink “6186-1”. We have added a white label “Digitally photographed by
L.M.Hernández 2005 right + left gill, cocoon”.]. Same data as the holotype- 3 larvae (on the same
slide), 1& (reared) [PARATYPES] [The slide with the larva has number “6186” and the female
“6186-2”].
OTHER MATERIAL
BRAZIL
Amapá State
PINNED
BR 156 Estrada Macapá-Oiapoque, Igarapé do Enrique (No. 3), 01º46´N50º52´W; 18.vi.2001,
(N. Hamada)—1% (without head and poor legs and wings) (INPA). BR 166, Estrada MacapáOiapoque, Igarapé Davi Grande, (No. 4), 01º 55’N 50º 51’W; 22.vi.2001, (N. Hamada)—2&&,
2%% (reared but not associated with pupae) (INPA).
SLIDE
BR 166, Estrada Macapá Oiapoque, Igarapé Davi Grande (No. 4), 01º55’N50º 51’ W; 22.vi.
2001, (N. Hamada)—3 pupae (INPA). BR 166, Estrada Macapá Oiapoque, Igarapé David Grande,
No. 4, 01º55’N 50º 51’ W; 24.vii. 2001, (N. Hamada)—1%, 2 pupal exuviae (INPA). Igarapé Agua
Branca, 02º40’N51º21’ W; 25.vii.2000, (N. Hamada)—3&&, several pupae (INPA). Fazenda
Monte Taboas, Rio Flechal, Braço do Araguari, 00º29’N51º8’W; 24.vii.2000, (N. Hamada)—1%,
pupa (slide 4) (INPA).
Simulium (Psaroniocompsa) delponteianum Wygodzinsky, 1961
TYPE MATERIAL
ARGENTINA
PINNED
Buenos Aires, La Granja, La Plata, Arroyo El Gato; 5.iii.1960, [Without collector’s name, but
Coscarón according to the original description of this species.]—1& (reared) only thorax, one wing,
two legs, and pupal exuviae (head, abdomen, one wing, one hind leg on slide), 2%% (reared)
[PARATYPES] (AMNH; NMUH).
54
SHELLEY ET AL.
ZOOTAXA
SLIDE
Buenos Aires, La Granja, La Plata, Arroyo El Gato; 5.iii.1960, [Without collectors’ names, but
they have been collected by Coscarón according to the original description of this species.]—1&
(reared) only head, abdomen, one wing, one hind leg (thorax, one wing, two legs, and pupal exuviae
pinned) [PARATYPE] (AMNH). Locality as paratype; 20.iii.1960, [Without collector’s name.]—
1&, 1% (reared), 2&&, 2%% (reared, but not associated with pupa), (AMNH).
OTHER MATERIAL
ARGENTINA
PINNED
Arroyo El Gato; 30.viii.1965, (Bach.)—1& (reared) (NMUH). Arroyo El Gato; 13.v.1966,
(Coscarón)—1% (reared) (NMUH). Bs. As., La Plata, La Granja; 20.iii.1960, [Without collector’s
name.]—1 larva (AMNH). Arroyo El Gato; 30.viii.1965, (Bach.) – 1& (reared) only thorax, one
wing, legs, and pupal exuviae (head, abdomen, and genitalia on slide) (NMUH). Arroyo El Gato;
13.v.1966, (Coscarón)—1% (reared) (NMUH).
SLIDE
Arroyo El Gato; 30.viii.1965, (Bach.)—1& (reared); only head, abdomen, and genitalia
(thorax, one wing, legs, and pupal exuviae pinned) (NMUH).
SPIRIT
Arroyo Carnaral; [Without date or collector’s name.]—several pupae and adults (BMNH) [The
specimens are in poor condition being almost transparent].
Simulium (Psaroniocompsa) ganalesense Vargas, Martínez Palácios & Díaz Nájera, 1946
All material examined listed by Shelley et al. (2002) was examined for this paper. Further material
is as follows:
HONDURAS
Gracias a Dios Department
SLIDE
Vila de Misawas, Patuca River; xi.1984, (G.B.White)—3&& (man-biting) (BMNH).
[These specimens may be S. sanguineum.]
Simulium (Psaroniocompsa) guaporense Py-Daniel, 1989
TYPE MATERIAL
BRAZIL
SIMULIUM AMAZONICUM
55
1274
ZOOTAXA
1274
Rondônia State
SLIDE
Estrada RO 399, Ig. Ponte de Pedra, Vilhena, RO; [Without date or collector’s name]—1 pupa
[HOLOTYPE, 5849-1], 1 larva [PARATYPE, 5849-2] (INPA). [The slide has a white label attached
to it by a transparent sticky tape and handwritten on one side of the slide in black ink “5849-1”. On
the side of the slide there are handwritten labels “HOLOTYPE, PARATYPE”. We have added a
white label in Luis Hernandez’ hand. “Digitally photographed by L.M.Hernández 2005”.]
OTHER MATERIAL
BRAZIL
Rondônia State
PINNED
Vilhena, Estrada Velha Colorado d´Oeste/ Nova Conquista, entrada a mais ou menos 23 km da
BR 364; 12º54’S 60º14’W; 11.viii.2002, (N. Hamada)—1%, 1& (reared, but not associated with
pupa) (INPA); 23.vii.2004, (N. Hamada)—1&, 1% without head (reared, associated with pupa)
(INPA). Vilhena, Estrada para Usina de Rio Vermelho, (no. 14), 12º54’S 60º14’W; 12.viii.2002, (N.
Hamada)—1&, 1% (reared, but not associated with pupae) (INPA); 25.vii.2004, (N. Hamada)—
3&&, 2%% (reared, associated with pupae) (INPA).
SLIDE
BR 364; 12º54’S 60º14’W; 11.viii.2002, (N. Hamada)—1&, 1% (reared), several pupae (BMNH;
INPA, laminas 1, 3-4); 25.vii.2004, (N. Hamada)—1&, biting humans (INPA).
SPIRIT
BR 364; 12º54’S 60º14’W; 11.viii.2002, (N. Hamada)—2 pupae (BMNH); 25.vii.2004, (N.
Hamada)—2 larvae (INPA).
Mato Grosso State
PINNED
Juina, Estrada Vilhena- Juina, BR 174, Roda d`água, (no. 10), 11º51’S 59º20’W; 6.viii.2002,
(N. Hamada)—2&& (reared, but not associated with pupa), 1% (reared) (INPA); 27.vii.2004, (N.
Hamada)—1& (reared, associated with pupa) (INPA). Comodoro, pequeno igarapé na Estrada para
PCH Cabixí II, ponte coberta com terra, 12º58’S 60º07’W, 26.vii.2004, (N. Hamada)—1&, 1%
(reared, associated with pupa) (INPA).
SLIDE
Juina, Estrada Vilhena—Juina, BR 174, Roda d`água, (no. 10), 11º51’S; 59º20’W; 6.viii.2002,
(N. Hamada)—1& (reared, but not associated with pupa) (INPA); 9&& (biting humans) (INPA).
Simulium (Psaroniocompsa) minusculum Lutz, 1910
56
SHELLEY ET AL.
All material listed under Shelley et al. (1982, 2001) was examined for this paper. Specimens that
have been subsequently imaged and to which new labels have been added, as well as further
material examined are as follows:
TYPE MATERIAL
BRAZIL
Minas Gerais State
PINNED
Lassance; iv-v.1910, (C. Chagas)—1& (IOC). [The specimen bears several labels: 1- Round
label with purple edge “Lectotype”; 2- White label “Simulium minusculum Lutz”; 3- White label
“Lectotype Simulium minusculum Lutz designated by A.J.Shelley & A.P.A. Luna Dias, 1981”; 4Syntype of S. minusculum Lutz examined A.J.Shelley 1980”; 5- White label with locality data
handwritten in black ink; 6- White label with date and collector’s name handwritten in black ink,
and “12.553”. We have added another label: 7- White label “Digitally photographed th. ant.+
posterior, abdomen L.M. Hernández 2006.”]
OTHER MATERIAL
BRAZIL
Goiás State
PINNED
Ceres, R. das Almas (site 813); 3.xi.1990, (A.J.Shelley & A.P.A.Luna Dias)—3&&, 2%%
(reared) (BMNH; IOC). Fazenda Luis Parrião, R.Mucambão; 19.iii.1995, (A.J.Shelley &
A.P.A.Luna Dias)—4&&, 6%% (reared) (BMNH; IOC). River Cana Brava; 7.1995, (M. Camargo)—
numerous && (man-biting). Near Aparecida do Rio Doce, Fazenda Tabatupã, Rio Doce (site 1443),
18º17’S 51º7’W 1851 ft; 4.viii.2001, (A.J.Shelley, M. Maia-Herzog & A.P.A.Luna Dias)—6&&
(man-biting) (IOC). Municipio Formoso, Rio Santa Teresa, (site 1150); 14.viii.1996, (A.J.Shelley,
A.P.A.Luna Dias & P.R. Garritano)—1& (man-biting) (IOC). Fazenda Margem Esquerda II, Rio
Cana Brava, (site 1155); 15.viii.1996, (A.J.Shelley, A.P.A.Luna Dias & P.R. Garritano)—3&&
(man-biting) (IOC). Rio Cristalino, Estrada que liga Minaçu a Formoso, (site 1176); 24.viii.1996,
(A.J.Shelley, A.P.A.Luna Dias & P.R. Garritano)—3&& (man-biting) (IOC). Rio cerca de
Niquelândia, (site 1111); 16.v.1996, (A.P.A.Luna Dias & P.R.Garritano)—1% (reared) (IOC).
SLIDE
Minaçu, R. Bonito; 16.iii.1995, (M. Camargo)—1& (BMNH). Minaçu, R. Bonito; 26.v.1995,
(C. Lowry & A.P.A.Luna Dias)—2&&, 2%% (BMNH).
Minas Gerais State
SLIDE
Lassance, Rio das Velhas, (site 580-3, 6, 8); 21-25.v.1981, (A.P.A.Luna Dias & P. Garritano)—
3&&, 2%% (one % reared), 4 pupae (BMNH).
Pará State
SIMULIUM AMAZONICUM
57
ZOOTAXA
1274
ZOOTAXA
1274
SLIDE
Near Altamira, Laranjal, R. Iriri, 52º53’W3º50’S; vii.1984, (T. Harvey)—1% (only genitalia,
rest of adult, and Pupal pelt pinned) (BMNH).
Tocantins State
PINNED
Conceição do Tocantins/Arraia, km 61 R. Arraia, (site 1235); 25.iv.1995, (P.A. Luna Dias &
P.R.Garritano)—2&&, 1% (reared) (BMNH). 5 km from Peixe, R. Tocantins, Corredeira da Bocão
(site 1001); 20.vi.1994, B.M.1994-254, (A.J.Shelley & A.P.A.Luna Dias)—4&&, 4%% (reared)
(BMNH; IOC). Pedro Alfonso, R. Sono, (site 987); 17.vi.1994, B.M.1994-254 (A.J.Shelley &
A.P.A.Luna Dias)—1& (reared) (BMNH). Monte Santo do Paraiso do Tocantins Rd, km 155 R.
Piedade, (site 987); 19.vi.1994, B.M.1994-254, (A.J.Shelley & A.P.A.Luna Dias)—1% (reared)
(BMNH). Nr. Natividade, R. Manuel Alves (site 1005); 20.vi.1994, B.M.1994-254, (A.J.Shelley &
A.P.A.Luna Dias) —1& (reared) (BMNH). Tocantinopolis/Novo Sitio, km 75, R. São Domingo,
(site 965); 11.vi.1994, B.M.1994-254, (A.J.Shelley & A.P.A.Luna Dias)—2&&, 2%% (reared)
(BMNH; IOC). Rio Chupe, (site 1395); 7.xi.1998, (A.J.Shelley, A.P.A.Luna Dias &
P.R.Garritano)—1& (man-biting) (BMNH). Itaituba, Ramal do km 11, Rio Itapacurá Grande
(cachoeira), (site 1387); 31.x.1998, (A.P.A. Luna Dias & P. R. Garritano)—1& (man-biting)
(BMNH). Peixe, 15 km de Baiana, TO 498, un-named stream, (EN 02), 12º17´09 0´´S, 48º 36´
09,8´´W; 3.ii.2004, (M.MH, APLD, MCA, VVAZ)—1 pupa (IOC). Peixe, road that connects Peixe
with São Salvador, (TO 280), un-named stream, (EN 20), 12º17´34,7´´S, 48º 23´57,9´´W; 4.ii.2004,
(M. M-H, A.P.L.D., M.C.A., V. V.A.Z.)—1& (man-biting) (IOC). Rio São João, Estrada que liga
Palmas a Porto Nacional, (site 1394); 7.xi.1998, (A.J.Shelley, A.P.A.Luna Dias & P.R. Garritano)—
1& (man-biting) (IOC). Rio Chupe, à 14 km do São João, (site 1395); 7.xi.1998, (A.J.Shelley,
A.P.A.Luna Dias & P.R. Garritano)—3 && (man-biting) (IOC). Monte Santo à 9 km de Paraíso,
Estrada Paraiso, Rio do Coco, (site 1227); 23.iv.1997, (A.P.A.Luna Dias)—1& (man-biting) (IOC).
Indo para Porto Nacional (26 km despois de Aueni), Rio a 45 km de Palmas, (site 1231);
24.ix.1997, (A.P.A.Luna Dias) (IOC).
Mato Grosso State
PINNED
Nr Rondonópolis, Rondonópolis/Guiratinga Rd, Fazenda Dona Ignacia, Tributary R.
Vermelho, R.Tadarimani (site 809); 28.ix.1990, (A.J.Shelley & A.P.A.Luna Dias)—2&& (manbiting) (BMNH; IOC). Comodoro, 1º depois da cidade, indo para Vilhena, Rio São Geraldo, (site
1363); 16.iv.1998, (A.P.A.Luna Dias & R. Pellegrini)—6&&, 2%% (reared) (IOC). P.N. Xingú, (site
1261); 27.vii.1997, (A.P.A.Luna Dias)—4 && (man-biting) (IOC).
ARGENTINA
Jujuy Province
PINNED
Route 34, Fraile Pintado to Libertador/San Martin to Tartagal, Rio de las Pedras; [Without
date.], (A.J.Shelley)—1& (reared, pupa on slide) (BMNH, BM 150347).
58
SHELLEY ET AL.
SLIDE
Route 34, Fraile Pintado to Libertador/San Martin to Tartagal, Rio de las Pedras; [Without
date.], (A.J.Shelley)—1 pupa (& on slide) (BMNH, BM 150347).
Salta Province
PINNED
Route 5, Oran to Las Lajitas, Rio del Valle; 29.i.1998, (A.J.Shelley)—1& (reared, pupa on
slide) (BMNH, BM 11998-16).
SLIDE
Route 5, Oran to Las Lajitas, Rio del Valle; 29.i.1998, (A.J.Shelley)—1 pupa (& on slide)
(BMNH, BM 11998-16).
Simulium (Psaroniocompsa) oyapockense Floch and Abonnenc (s.l.)
All material listed by Shelley et al. (1997) was examined for this paper and specimens listed as
oyapockense/roraimense are here regarded as oyapockense s.l. Further material examined is as
follows:
BRAZIL
Amapá State
PINNED
Macapá-Oiapoque Rd., Cachoeira do Rio, Amapá Grande, (site 1319) (9); 10.xii.1994, (S. Luz
& A.P.A.Luna Dias)—1& (reared) (BMNH). Municipio do Calçoene, Macapá-Oiapoque Rd, ASA
Aberta, R. Calçoene (cachoeira), (site 1320)(10); 11.xii.1994, (S. Luz & A.P.A.Luna Dias)—2&f,
#1% (reared) (BMNH). Oiapoque, Cachoeira Maripá, Rio Oiapoque; 13-16.xii.1994, (S.Luz &
A.P.A.Luna Dias)—1& (reared); only thorax (head, abdomen, wings, legs, genitalia, and pupal
exuviae on slide) (BMNH).
SLIDE
Oiapoque, Cachoeira Maripá, Rio Oiapoque; 13-16.xii.1994, (S.Luz & A.P.A.Luna Dias)—1&
(reared); only head, abdomen, wings, legs, genitalia, and pupal exuviae (thorax pinned) (BMNH).
Amazonas State
PINNED
Lábrea, Rio Mari, Estrada Lábrea-Humaitá (Transamazônica), a cerca de 30 km depois de
Lábrea, (site 1412); 9.viii.1999, (A.J. Shelley & A.P.A. Luna Dias)—1& (reared) (IOC). Igarapé
Tiquié, trib. of R. Vaupés; 15.xii.1977, (C. Vicente)—1& (man-biting) (BMNH).
Mato Grosso State
PINNED
SIMULIUM AMAZONICUM
59
ZOOTAXA
1274
ZOOTAXA
1274
Cuiabá, Municipio Cuiabá, Rd to Sto. Antonio from Cuiabá, Rio Cuiabá, (site 1512, L18),
15º54’76’’S 56º00’86’’W, 584ft; 21.x.2002, (A.J.Shelley, M.M.Herzog, A.P.A.Luna Dias &
L.M.Hernández)—1& (man-biting) (BMNH, B.M. 2002-187).
Mato Grosso do Sul State
PINNED
Mato Grosso do Sul State, Municipio Aquidauana, Aquidauana, at Fisherman’s Club, Rio
Aquidauana, (site 1528), 20º27’03’’S 55º49’7’’W, 512ft; 31.x.2002, (L.M.Hernández &
M.M.Herzog)—12&& (man-biting) (BMNH, BM 2002-187). Rd from Bonito to Aquidauana,
bridge at the border of Municipios Anastacio and Bonito, under bridge on the side of Municipio
Anastacio, Rio Miranda, (site 1526, L35), 20º45’78’’S 56º05’40’’W, 6155ft; 30.x.2002,
(L.M.Hernández)—15&& (man-biting) (BMNH, B.M. 2002-187; IOC). Municipio Aquidauana,
Aquidauana, at Fisherman’s Club, Rio Aquidauana, (site 1528, L37), 20º27’03’S 55º49’7W, 512ft;
31.x.2002, (L.M.Hernández & M.M.Herzog)—14&& (man-biting) (BMNH, B.M. 2002-187; IOC).
Minas Gerais State
PINNED
Sta. Rita do Sapucaí, Rio Sapucaí; 25.i.1983, (A.P.A.Luna Dias)—4&& (man-biting) (BMNH).
Santa Rita do Sapucaí, Rio Sapucaí, (site 641); 25.i.1983, 19.ii.1973, (A.P.A.Luna Dias)—13&&
(man-biting) (IOC; BMNH).
Paraná State
PINNED
São Salvador/Paranã, right margin of Paranã, Córrego Lages, (EN 39); 5.ii.2004, (M.M.H.,
A.P.L.D., M.C.A., V.V.A.Z.)—2&& (man-biting) (IOC).
Roraima State
PINNED
Rio Branco; 10.vii.87, (P. Horton)—17&& (man-biting) (BMNH; IOC). Uiramutão, Córrego
Caxirimã (site 1300); 29.x.1997, (A.J.Shelley & A.P.A.Luna Dias)—2&& (man-biting) (BMNH).
Vila Pereira, R. Surumu (rapids); 11.viii.1984, (A.J.Shelley & A.P.A.Luna Dias)—20&& (manbiting) (BMNH). Igarapé Murupu (site 955); 6.vi.1994, (A.P.A.Luna Dias)—1% (reared) (BMNH).
Rio Tacutu (site 1315); 28.xi.1994, (S. Luz & A.P.A.Luna Dias)—14&& (man-biting) (BMNH;
IOC), 1& (reared) (BMNH), 6%% (reared) (BMNH; IOC). Boa Vista-Caracarai Rd, R. Branco,
Cachoeira Bem Querer (site 952); 2.v.1994, (A.P.A.Luna Dias)—5&&, 4%% (reared) (BMNH;
IOC). Municipio de Caracarai, R. Branco, Cachoeira Bem Querer, (site 1313) (3); 24.xi.1994, (S.
Luz & A.P.A.Luna Dias)—8&& (man biting), 7&&, 6%% (reared) (BMNH; IOC). Bonfim, Rio
Tacutu, Cachoeira de Maloca do Pium (site 1316)(6); 29.xi.1994, (S. Luz & A.P.A.Luna Dias)—
12&& (man-biting) (BMNH; IOC), 1& (reared) (BMNH). Vila Pereira, Fazenda do Sr. Paulo; Cesar,
R. Surumu (cachoeira abaixo da ponte), (site 1318) (8); 30.xi.1994, (S. Luz & A.P.A.Luna Dias)—6
&& (man-biting) (BMNH; IOC).
SPIRIT
60
SHELLEY ET AL.
Nr Uiramutã, R. Maú, (site 1299); 20.x.1997, (A.J.Shelley & A.P.A.Luna Dias)—1 & (manbiting). Uiramutã, Igarapé, R. Ailã, (site 1297); 20.viii.1997 (A.J.Shelley & A.P.A.Luna Dias)—
numerous && (man-biting) (BMNH). Igarapé, Uiramutã, Cachoeira de Urucá, (site 1303);
29.x.1997, (A.J.Shelley & A.P.A.Luna Dias)—numerous && (man-biting) (BMNH). Nr. Uiramutã,
Maloca Canapan, (site 1301); 13.x.1997, (A.J.Shelley & A.P.A.Luna Dias)—numerous && (manbiting) (BMNH). Bem Querer, (site 786); 4.xii.1986, (A.J.Shelley & A.P.A.Luna Dias)—1 larva
(BMNH). Cachoeira Bem Querer, Rio Branco, Caracarai; 24.i.2003, (C. Rios Velásquez) —4&&,
6%% (reared) (BMNH).
Rio Grande do Sul State
SPIRIT
Near Irai, Rio Uruguai, nr frontier with Santa Catarina Province (BR 158), (site 876); 22.viii.1982,
(A.J.Shelley)—6&& (man-biting) (BMNH).
Tocantins State
PINNED
Araguatins to Cachoerinha-Wonderlandia, Rd, km 53, Rio São Martinho (Site 968); 12.vi.1994
(A.J.Shelley, M.Herzog & A.P.A.Luna Dias)—1&1% (reared) (BMNH). Peixe, 15 km de Baiana, TO
498, un-named stream, (EN 02), 12º 17´ S 48º36´09 ´´W; 3.ii.2004, (M.M.H., A.P.L.D., M.C.A. &
V.V.A.Z.)—1% (reared) (IOC). Peixe, road that connects Peixe with São Salvador, (TO 280), unnamed stream, (EN 20), 12º17´34,7´´S 48º23´57,9´´W; 4.ii.2004, (M.M.H., A.P.L.D., M.C.A. &
V.V.A.Z.)—1& (man-biting) (IOC).
COLOMBIA
PINNED
Comis. Vaupés, Caño Cuduyari, near Mitú; 18.i.1978, (M.A.Tidwell)—2&& (biting man)
(BMNH).
GUYANA
SPIRIT
Nr. Uiramutã [Brazil, R. Mau]; 31.x.1997, (A.J.Shelley & A.P.A.Luna Dias)—numerous
(man biting) (BMNH).
&&
PARAGUAY
SLIDE
Mbaracayú, Canindeyu River Jejui-mí, between Corona Bridge and the western border of the
reserve; 18.vii.1996, (C. Nouzille & L.Lagache)—1 pupal exuviae (adult in alcohol) (BMNH, B.M.
150617). Mbaracayú Forest Reserve, Chorro Morote; 14.vi.1995, (A.J.Shelley)—1& (reared, pupa
on slide) (BMNH, B.M. 1995-201).
SPIRIT
SIMULIUM AMAZONICUM
61
ZOOTAXA
1274
ZOOTAXA
1274
Mbaracayú, Canindeyu River Jejui-mí, between Corona Bridge and the western border of the
reserve, (site 37); 18.vii.1996, (C. Nouzille & L.Lagache)—1& (reared, pupa on slide) (BMNH,
B.M. 150617). Mbaracayú Forest Reserve, Chorro Morote, (site 7); 14.vi.1995, (A.J.Shelley)—1&
(reared, pupa on slide) (BMNH, BMNH (E) 15041b, B.M. 1995-201).
URUGUAY
SPIRIT
Salto Termas del Dayman, Rio Dayman; 23.ix.19992, (A.J.Shelley)—several && (man-biting)
(BMNH).
As S. sanchezi Ramírez Pérez, Yarzábal & Peterson
VENEZUELA
PINNED
Territorio Federal de Amazonas, Siquita; 31.vi.1982, [Without collector’s name.]—3&& (2&&
not associated with pupa; 1 & pupa on slide), 1% (reared, pupa on slide) (BMNH).
SLIDE
Same date as pinned adults—2 pupae (&, % pinned) (BMNH).
Simulium (Psaroniocompsa) roraimense Nunes de Mello, 1974
All material listed by Shelley et al. (1997) was examined for this paper. Further material examined
is as follows:
TYPE MATERIAL
BRAZIL
Roraima State
PINNED
[Without country, date or collector’s name, but agrees with the locality given in the original
description Nunes de Mello, 1974:45.]—1% (reared, not associated with pupa). [HOLOTYPE]
(INPA) Roraima territory [=State], Cachoeira, Rio Cauamé, 3 km above bridge on BR 174; xi.1972,
(J.A.Nunes de Mello). [The specimen bear several labels: 1- White, handwritten label “1001-1”; 2White, printed label “UA.ICB”; 3- White, handwritten, black-bordered label “Simulium roraimense
sp. n.”; 4- White, printed label “5881-1”; 5- Red label “Holotipo S. roraimense Mello, 1971. We
have added another label: 6- White label “Digitally photographed by L.M.Hernández 2005”.]
OTHER MATERIAL
BRAZIL
Roraima State
62
SHELLEY ET AL.
ZOOTAXA
PINNED
Near Uiramutã, R. Ailã, (site 1297); 28.x.1997, (A.J.Shelley & A.P.A.Luna Dias)—4&&, 2%%
(reared) (BMNH; IOC). Uiramutã/Normandia Rd, R. Maú, (site 1299); 19.x.1997, (A.J.Shelley &
A.P.A.Luna Dias)—7&&, 8%% (reared) (BMNH; IOC). Nr. Boa Vista, Igarapé Murupu (site 956);
6.v.1994, (A.P.A.Luna Dias)—1&, 2%% (reared), 1& (man-biting) (BMNH). R.Cauamé
(Cachoeira), (site 1311); 21.xi.1994, (S. Luz & A.P.A.Luna Dias)—2&&, 4%% (BMNH; IOC).
Cachoeira, R. Cauamé; 12.vii.1984, (A.J.Shelley & A.P.A.Luna Dias)—10&& (man-biting), 1%
(reared) (BMNH; IOC). Cachoeira, R. Cauamé, (site 539); 22.xi.1980, (A.J.Shelley)—1& (reared)
only thorax and pupal exuviae in glycerine (head, abdomen, two wings, genitalia, and four legs on
slide), 1% (reared) only thorax and pupal exuviae in glycerine (abdomen, one wing, and four legs on
slide) (BMNH).
SLIDE
Mission Post, R.Catrimani; 9.i.1977, (A.J.Shelley)—9&&, 5%% (reared) (BMNH). Cachoeira,
R. Cauamé, (site 539); 22.xi.1980, (A.J.Shelley)—1& (reared) only head, abdomen, two wings,
genitalia and four legs (thorax, and pupal exuviae pinned); 1% (reared) only abdomen, one wing,
and four legs; thorax pinned and pupal exuviae in glycerine) (BMNH).
SPIRIT
Near Boa Vista, R.Cauamé, 12.iv.1979, (A.J.Shelley)—1& (man-biting) (BMNH).
ECUADOR
Manabi Province
PINNED
Porto Viejo, Pocahonda Rd, Pachinche, Rio Porto Viejo, (site 142); 6.xii.1984, (M. Arzube)—
3&&, 6%% (reared, 3&&, 4%% with pupa on slide) (BMNH).
SLIDE
Porto Viejo, Pocahonda Rd, Pachinche, Rio Porto Viejo, (site 142); 6.xii.1984, (M. Arzube)—7
pupae (3&&, 4%%; adults pinned) (BMNH).
Simulium (Psaroniocompsa) sanguineum Knab, 1915
TYPE MATERIAL
COLOMBIA
PINNED
Boca de Arquia, Atrato River; [Without date or collector’s name.]—& [HOLOTYPE] (BMNH)
[The specimen bears several labels: 1- White, red-rounded label “Holotype”; 2- Small, orange label
“Type”; 3-White label with locality information; 4- White label “Simulium sanguineum Type
Knab”; 3- White label “ Digital Photograph BMNH (E)#625035”; 5- White label “Digital
photographs th. anter. + post., abdomen by L.M.Hernández BMNH 2004”. Same locality as the
holotype- 1& [PARATYPE] (BMNH).
SIMULIUM AMAZONICUM
63
1274
ZOOTAXA
1274
OTHER MATERIAL
COLOMBIA
Department of Cauca
PINNED
Approx. 11 km n. de Santander de Quilachao, Finca da Cuprecia, (site 78-26, 320-1, other as
slide material); 11.i.1977, (M.A.Tidwell)—1&, 1% (not associated with pupae).
SLIDE
La Cuprecia, approx. 11 km n. de Santander de Quilachao; 11.i.1977, (M.A.Tidwell)—1& (not
associated with pupa), 1 pupa (adult pinned, no. 320-1) (BMNH).
Department of Chocó
PINNED
Near Tagachi, 1 km from Boca Arquia, Nr. Rio Atrato, (site 78.28b, 77.3); 9.xii.1978,
(M.A.Tidwell)—4&&, 3%% (some specimens with pupal exuviae on slides; 1& only thorax (head,
wings, legs, abdomen, and genitalia on slide) (BMNH).
SLIDE
Near Tagachi, approx. 1 km from Boca Arquia, (site 78.28b, 77.3); 9.xii.1978, (M.A.Tidwell)—
4&&, 2%% (not associated with pupae), 1& only head, wings, legs, abdomen and genitalia (thorax
pinned), several pupal exuviae (BMNH). R. Tagachi, nr. R. Atrato, approx. [1km] from Boca
Arquia, (site 403-1-8); 12.ix.1978, (M.A.Tidwell)—3&&, 4%% (not associated with pupae), several
pupae (BMNH).
PANAMA
Darien Province
PINNED
Tucuti, (no. 1); 29.i.1933, (L.H.Dunn)—2&& (man-biting) (BMNH). Boca de Tuquasa, R.
Chucunaque, (no. 26); 7.ii.1933, (L.H.Dunn)—7&& (BMNH).
SLIDE
Canal Zone, Darien, Boca Metiti, Rio Chucunaque; 18.i.1995, (J.Petersen & J.E.Canon)—2
larvae (BMNH).
Simulium (Psaroniocompsa) siolii Py-Daniel, 1988
TYPE MATERIAL
As Simulium (Psaroniocompsa) lourencoi Py-Daniel, 1988 [Synonymised with S. siolii in this
paper.]
64
SHELLEY ET AL.
BRAZIL
Goiás State
ZOOTAXA
1274
SLIDE
Rio Bandeira, Br. 153, Entre Araguaiana e Imperatriz; 4.iii.1987, (Py-Daniel)—1 pupal
exuviae [HOLOTYPE, INPA 6190], 3 pupae [PARATYPES, INPA 6190-2-4) (INPA) [The
holotype is mounted together with three paratypes. On one side of the slide each specimen has a
number together with its type status (“Holotipo” or “Paratipo”). On the other side of the slide there
is a white, red-bordered label with the printed locality information, date and collector’s name. It
also reads “Shelleyellum lourencoi INPA 6190”. On the left hand side corner there is a number,
handwritten in black pen “320, 18”. At the lower, right hand side corner “6190”. We have added
another label “Digitally photographed by L.M.Hernández 2005 Pupa-paratype-2”.]. Same date as
holotype- 1& and pupal exuviae [PARATYPE, INPA 6190-5] (INPA) [The slide has on one side
handwritten “6190-5 PARATIPO (18) S. lourencoi 6190”. On the other side there is a white label
with locality information, date and collector’s name printed on the label. It also reads “Shelleyellum
lourencoi”.]
OTHER MATERIAL
BRAZIL
Goiás State
PINNED
Formoso, Rio Pipoca, (site 1368); 10.v.1998, (A.J.Shelley, M.M.Herzog & A.P.A.Luna Dias)—
3&&, 1% (reared) (IOC).
Maranhão State
PINNED
Previously identified as Simulium (Psaroniocompsa) lourencoi Py-Daniel: Cipo, afluente do
Itapecuru, 07º 24’ S 47º12’ W; 12.viii.2001, (N. Hamada)—1% (reared but not associated with
pupa, genitalia and one wing on slide) (INPA). Igarapé Itapecuru, Cachoeira Itapecuruzinho
(balneário), 07º24’S47º12’W; 8.viii.2001, (N. Hamada)—1& (reared, but not associated with pupa;
head, wings, legs, abdomen, and genitalia on slides) (INPA).
SLIDE
Previously identified as Simulium (Psaroniocompsa) lourencoi Py-Daniel: Cipo, afluente do
Itapecuru, 07º 24’ S 47º 12’ W; 12.viii.2001, (N. Hamada)—1% (reared but not associated with
pupa, only genitalia and one wing, rest of adult pinned) (INPA). Igarapé Itapecuru, Cachoeira
Itapecuruzinho (balneário), 07º24’S 47º12’W; 8.viii.2001, (N. Hamada)—1& (reared, but not
associated with pupa; only head, wings, legs, abdomen, and genitalia; rest of adult on pin) (INPA).
Carolina, Igarapé Itapecuru, Cachoeira Itapecuruzinho (balneário), 07º24’S 47º12’W; 8.viii.2001,
(N. Hamada)—2&&, 1% (reared) (INPA, laminas 1-2, 4). Same as previous locality; 12.viii.2001,
(N. Hamada)—1& (without pupa) (INPA, lámina 3). Carolina, Represa Itapecuru; 25.viii.2002, (N.
Hamada)—3 pupae (INPA; BMNH).
SPIRIT
SIMULIUM AMAZONICUM
65
ZOOTAXA
1274
Previously identified as Simulium (Psaroniocompsa) lourencoi Py-Daniel: Carolina, Represa
Itapecuru; 25.viii.2002, (N. Hamada)—several pupae (INPA).
Mato Grosso State
PINNED
Sorriso/Sinop Rd, R. Celeste, (site 1241); 21.vi.1997, (A.P.A. Luna Dias & P.R.Garritano) —
1% (reared) (BMNH). Tangará da Serra, stream 7 km from Riberão do Sapo, Parecis to Tangará,
Fazenda Santa Carmen, (site 1049); 23.v.1995, (A.P.A. Luna Dias & P.R.Garritano, M.M.Elazaro,
M. Leila)—10 (man-biting), 4&&, 6%%(reared) (IOC; BMNH). Sorriso, Rio Lira, (site 1240);
21.vi.1997, (A.P.A. Luna Dias)—3&& (man-biting) (IOC; BMNH). Estrada que liga Sorriso a
Sinop, (site 1241); 21.vi.1997, (A.P.A.Luna Dias)—11&& (man-biting) (IOC; BMNH). Estrada
Arenápolis—Parecis, 8,3 Km de Marilândia, coleta no Rio Sucuri, (site 1057); 18.v.1995,
(A.P.A.Luna Dias, P.G.Garritano, M.M.Elázaro, M.Leila)—1&, 1% (reared) (IOC). Rio Ávila, a 50
km de Vilhela (site 1364); 16.iv.1998, (A.P.A. Luna Dias & R. Pellegrini)—5&& (man-biting)
(IOC).
SLIDE
Sorriso/Sinop Rd, R. Celeste, (site 1241); 21.vi.1997, (A.P.A.Luna Dias & P.R.Garritano) —1
pupa (& pinned) (BMNH).
Mato Grosso do Sul State
PINNED
Itiquira, Cachoeira BR 163, (site 666); 11.ii.1984, (A.P.A. Luna Dias)—1&, 1% (reared)
(BMNH). Alto Araguaia-Alto Taquari-Coxim Rd, stream between Buritis and Pedro Gomes, (site
1444), 17º49’S 53º49’W, 1796 ft; 6.viii.2001, (A.J.Shelley, M. Maia-Herzog & A.P.A.Luna Dias)—
1& (reared) (BMNH).
SLIDE
Cach. BR 163, (site 663-5); 11.ii.1984, (A.P.A.Luna Dias)—2 pupal exuviae (&, % pinned)
(BMNH). Alto Araguaia-Alto Taquari-Coxim Rd, stream between Buritis and Pedro Gomes, (site
1444), 17º49’S 53º49’W, 1796 ft; 6.viii.2001, (A.J.Shelley, M. Maia-Herzog & A.P.A.Luna Dias)—
1 pupa (& pinned) (BMNH).
Tocantins State
PINNED
Pedro Alfonso/Tocantins Rd, km 86 Córrego Piabanha, (site 989); 17.vi.1994, (A.J.Shelley &
A.P.A.Luna Dias)—1& (reared) (BMNH, B.M.-1994-254). Abreulandia/ Divinópolis Rd, 12 km
before state border, córrego, (site 997); 18.vi.1997, (A.J.Shelley & A.P.A.Luna Dias)—1&, 1%
(reared) (BMNH, B.M.1994-254; IOC). Tocantinópolis to Araguatins Rd, Riberão Grande, (site
963); 11.vi.1994, (A.J.Shelley & A.P.A.Luna Dias)—5&&, 1% (reared) (BMNH, B.M.1994-254).
Macaúba/São Bento Rd, km 53, R. São Martinho, (site 968); 12.vi.1994, (A.J.Shelley & A.P.A.Luna
Dias)—1& (reared) (BMNH, B.M.1994-254). Tocantinópolis to Araguatins Rd, km 31, R. Butica,
(site 964); 11.vi.1994, (A.J.Shelley & A.P.A.Luna Dias)—2&&, 2%% (reared) (BMNH, B.M.1994-
66
SHELLEY ET AL.
254; IOC). Rio a 45 Km de Palmas, indo para Porto Nacional, 26 Km depois de Aureni 3, (site
1231); 24.iv.1997, (A.P.A.Luna Dias)—1& (man-biting) (BMNH). Rio Araguaia, (site 978);
14.vi.1994, (A.J. Shelley, Maia- Herzog, A.P.A.Luna Dias)—1% (reared) (IOC). Peixe, 15 km de
Baiana, TO 498, un-named stream, (EN 02), 12º17´09,0´´S, 48º36´09,8´´W; 3.ii.2004, (M.M.H.,
A.P.L.D., M.C.A., V.V.A.Z.)—2&& (reared), 5&& (man-biting) (IOC).
SLIDE
Tocantinopolis to Araguatins Rd, Riberão Grande, (site 963); 11.vi.1994, (A.J.Shelley, M.
Herzog & A.P.A.Luna Dias)—7 pupae (adult pinned), 2%% (reared) (some specimens with BMNH,
B.M.1994-254). Tocantinopolis to Araguatins Rd, Km 32, Rio Butica, (site 964); 11.vi.1994,
(A.J.Shelley, M. Herzog & A.P.A.Luna Dias)—1 pupa (adult pinned), 1% (reared), 3&& (manbiting) (BMNH). Araguatins to Cachoeirinha Wonderlandia Rd., km 53, Rio São Martinho, (site
968); 12.vi.1994, (A.J.Shelley, M.Herzog & A.P.A.Luna Dias)—1 pupa (adult reared). BelémBrasília Rd, BR 153, Pedro Alfonso to Tocantins Rd, km 150 before Miracema, Rio Vermelho, (site
989); 17.vi.1994 (A.J. Shelley, M. Herzog & A.P.A.Luna Dias)—1 pupa (& pinned) (BMNH).
Abreulandia/Divinópolis Rd, 12 km before state border, córrego, (site 997); 18.vi.1997,
(A.J.Shelley & A.P.A.Luna Dias)—1 pupa (& pinned) (BMNH).
SPIRIT
Tocantinópolis to Araguatins Rd, Riberão Grande, (site 963); 11.vi.1994, (A.J.Shelley, M.
Herzog & A.P.A.Luna Dias)—1& (man-biting), 1&1% (reared, pupae on slide), many pupae, 3
larvae (BMNH). Tocantinopolis to Araguatins Rd, Km 32, Rio Butica, (site 964); 11.vi.1994,
(A.J.Shelley, M. Herzog & A.P.A.Luna Dias)—3&& (man-biting), 3 pupae (BMNH). Araguatins to
Cachoerinha-Wonderlandia Rd, km 53, Rio São Martinho, (site 968); 12.vi.1994 (A.J.Shelley,
M.Herzog & A.P.A.Luna Dias)—numerous larvae and pupae (BMNH). Belém-Brasília Rd, BR 153,
Pedro Alfonso to Tocantins Rd, km 150 before Miracema, Rio Vermelho, (site 989); 17.vi.1994
(A.J. Shelley, M.Herzog & A.P.A.Luna Dias)—many larvae, pupae (BMNH).
Simulium (Psaroniocompsa) tergospinosum Hamada, 2000
TYPE MATERIAL
BRAZIL
Amazonas State
PINNED
Apuí, Cachoeira Apuí, Rio Juma, 07º11´S59º54´W; 1-2.viii.1998, (F.F.Xavier Filho)—1%
(reared) [HOLOTYPE] (INPA) [The holotype bear several labels: 1- Red label “Holótipo”; 2White label with locality data and collector’s name; 3-White label in Hamada’s hands “Simulium
tergospinosum Hamada, 2000”. We have added another label: 4- White label “Digitally
photographed parts Thoracic pattern (ant+post.), pupa in alcohol, abdomen. Taken by
L.M.Hernández (BMNH 2004)”. Same locality as holotype- 3&&, 2%% (reared, but not associated
with pupa) [PARATYPES] (INPA).
SPIRIT
Apuí, Cachoeira Apuí, Rio Juma, 07º 11´S 59º 54´W; 3.vii.1998, (N. Hamada)—several pupae
SIMULIUM AMAZONICUM
67
ZOOTAXA
1274
ZOOTAXA
1274
and larvae [PARATYPES] (BMNH).
OTHER MATERIAL
SLIDE
Apuí, Cachoeira Apuí, Rio Juma, 07º11´S 59º54´W; 1-2.viii.1998, (F.F.Xavier Filho)—2 pupae
(INPA, lamina 1). Apuí, Cachoeira Apuí, Rio Juma, 07º11´S59º54´W; viii.1998, (F.F.Xavier
Filho)—4&&, 2%% (reared, but not associated with pupae), several pupae (INPA, laminas 2-7).
Apuí, Igarapé Seringueira; 2.vii.1998, (N. Hamada)—several pupae (INPA). Apuí , Cachoeira
Apuí, Rio Juma; 3.v.1999, (N. Hamada)—2 pupae (BMNH).
SPIRIT
Apuí, Cachoeira Apuí, Rio Juma, 07º 11´S 59º 54´W; 3.v.1999, (N. Hamada)—several pupae
(INPA).
Simulium (Psaroniocompsa) venezuelense Ramírez Pérez & Peterson, 1981
BRAZIL
Amapá State
PINNED
R. Patanari (trib. of R. Oyapock); 22.v.1992, (C. Lowry & A.P.A.Luna Dias)—1% (reared)
(BMNH). Oiapoque, R. Patanari, Cachoeira Patanari, (site 1322); 14.xii.1994, (S.Luz &
A.P.A.LunaDias)—1&, 1% (reared, pupa on slides) (BMNH; IOC). Oyapoque, Km 9 da Oiapoque/
Macapá Rd, R. Patanari, Cachoeira Patanari II (site 1323); 14.xii.1994, (S.Luz & A.P.A.Luna
Dias)—4&&, 3%% (reared, pupae on slides) (BMNH; IOC).
SLIDE
Oiapoque, R. Patanari, Cachoeira Patanari, (site 1322); 14.xii.1994, (S. Luz & A.P.A.Luna
Dias)—2 pupal exuviae (&, % pinned), 1% (reared) (BMNH; IOC). Oiapoque, Km 9 da Oiapoque/
Macapá Rd, R. Patanari, Cachoeira Patanari II (site 1323); 14.xii.1994, (S.Luz & A.P.A.Luna
Dias)—4 pupal exuviae (&, % adults pinned), 1&, 1% (reared), 1 pupa (adult lost) (BMNH; IOC).
Oiapoque, R. Patanari, Cachoeira Patanari; 11-12.vii.2002, (S. Luz)—%% (reared) (FioCruz,
Manaus; BMNH) [adults sent to Sergio Lutz for DNA studies in FioCruz, Manaus, genitalia and
pupae on slides at BMNH]. Oiapoque, R. Patanari, Cachoeira Patanari, (site 1322); 14.xii.1994, (S.
Luz & A.P.A.Luna Dias)—1% (reared) (BMNH).
68
SHELLEY ET AL.

Morphological overview of the amazonicum

Transcription

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