Chironomidae (Insecta: Diptera) of Ecuadorian Highaltitude Streams: A Survey and Illustrated Key
Chironomidae (Insecta: Diptera) of Ecuadorian Highaltitude Streams: A Survey and Illustrated Key
Chironomidae (Insecta: Diptera) of Ecuadorian Highaltitude Streams: A Survey and Illustrated Key
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Abstract
Chironomidae (Diptera) are among the most diverse and widespread aquatic insects, with roughly 5,500 described species inhabiting an enormous
variety of aquatic ecosystems, ranging from moist soils to lakes and rivers, and even marine ecosystems. Despite its ubiquity, the group remains
underrepresented in studies addressing aquatic insect assemblages of high-altitude systems, either glacier-fed or non-glacial, particularly in South
America. Glacier-fed streams possibly are one of the harshest ecosystems in nature, and present a distinct downstream pattern in species distribution
and diversity away from the constraining influence of the glacier. In this context, the goal of our study was to provide data on the chironomid fauna
of glacier-fed streams, together with neighboring non-glacial streams in Ecuador, in order to contribute to the overall knowledge of tropical fauna in
high Andean regions. Also, we sought to provide an identification key and photography material for future studies. Collections of non-biting midges
were made in Ecuador during Jan 2008. In total, 1,412 specimens belonging to 16 genera and at least 23 species within the subfamilies Chironominae
(3 taxa), Diamesinae (3 taxa), Podonominae (3 taxa), Orthocladiinae (13 taxa), and Tanypodinae (1 taxon) were found. The subfamilies Podonominae
and Orthocladiinae predominated in terms of abundance. Parochlus was the most widespread genus of Chironomidae, while Cricotopus was the
most diverse. This study contributes to the knowledge of the chironomid fauna in the high-altitude streams in Ecuador, and hopefully will motivate
further studies in the area.
Key Words: non-biting midges; high Andes; Neotropical; Antisana volcano; glacier-fed streams; non-glacial streams
Resumo
Chironomidae (Diptera) estão entre os mais diversos e difundidos insetos aquáticos, com cerca de 5.500 espécies descritas, habitando uma enorme
variedade de ecossistemas aquáticos, desde solos úmidos, lagos e rios, até mesmo ecossistemas marinhos. Apesar de sua onipresença, o grupo
continua sub-representado em estudos que abordam assembléias de insetos aquáticos de sistemas de alta altitude, seja de origem glacial ou não,
particularmente na América do Sul. Riachos glaciais são possivelmente um dos ecossistemas mais severos da natureza e, portanto, apresentam tipi-
camente um padrão espacial distinto na distribuição e diversidade das assembleias de espécies a jusante e longe da influência limitadora da geleira.
Neste contexto, o objetivo do nosso estudo foi fornecer dados sobre a fauna de Chironomidae de alguns riachos glaciais e não-glaciais no Equador,
a fim de contribuir para o conhecimento global da fauna tropical nas regiões andinas de alta latitude, além de fornecer uma chave de identificação e
um catálogo fotos para futuros estudos. Durante o mês de janeiro de 2008, as coletas de Chironomidae foram realizadas no Equador. No total, 1.412
espécimes pertencentes a 16 gêneros e pelo menos 23 espécies pertencentes às subfamílias Chironominae (3 táxones), Diamesinae (3 táxones),
Podonominae (3 táxones), Orthocladiinae (13 táxones) e Tanypodinae (1 taxon) foram encontrados. As subfamílias Podonominae e Orthocladiinae
predominaram em termos de abundância. Parochlus foi o gênero mais difundido, enquanto Cricotopus o mais diversificado. O presente estudo
contribui para o conhecimento sobre a fauna de Chironomidae nos córregos de alta altitude no Equador e espera-se que motive estudos adicionais
na área.
Palavras Chave: quironomídeos; Alto Andes; Neotropical; vulcão Antisana; riachos glaciais e não glaciais
Non-biting midges (Insecta: Diptera: Chironomidae) are true flies, gations (Silva & Ekrem 2016). Nevertheless, in order to obtain the most
and the most widely distributed free-living holometabolous insects (Fer- biologically informative data, it is fundamental to identify taxa to species,
rington 2008). Chironomids inhabit an enormous variety of aquatic eco- because within a single genus, species may respond in a distinct manner
systems, ranging from moist soils to pools in tree-holes, and from low- to environmental changes (Lenat & Resh 2001). Generally, the lack of
oxygen lake sediments to fast-flowing mountain streams (Ferrington et descriptions and keys to a local fauna precludes species determination,
al. 2008). The adult life stage of chironomids is ephemeral, and most of or workers choose to overlook the chironomids in favor of groups (e.g.,
the lifespan is spent in the larval stage (Thienemann 1954; Tokeshi 1995). Ephemeroptera, Plecoptera, Trichoptera) that are more limited in num-
The countless species and habitat diversity makes this family a valuable ber and diversity (Spies et al. 2009).
indicator species for lentic and lotic aquatic ecosystems, but also they Approximately 900 chironomid species are recognized from the
are particularly well suited for phylogenetic and biogeographical investi- Neotropical region (Martin Spies, personal communication). This
1
Polish Academy of Sciences, Institute of Geological Sciences, Warsaw, Poland; E-mail: ladislav.hamerlik@gmail.com (L. H.)
2
Matej Bel University, Department of Biology and Ecology, Banská Bystrica, Slovakia
3
University of São Paulo, Institute of Biosciences, Department of Zoology, Laboratory of Systematic and Biogeography of Insecta, São Paulo, São Paulo, Brazil;
E-mail: fabiologia@gmail.com (F. L. S.)
4
University of Copenhagen, Department of Biology, Freshwater Biological Section, Copenhagen, Denmark; E-mail: djacobsen@bio.ku.dk (D. J.)
*Corresponding author; E-mail: ladislav.hamerlik@gmail.com
Fig. 1. Map of the Antisana stream system. For site codes see Tables 1 and 2. Glaciated area is marked with grey. Sites 1 to 10 are glaciated, sites 11 to 17 are
non-glacial.
4. Prementum with 3 dense brushes of hair-like processes. Usually with distinctive ‘collar’ (occipital margin of the head). Mentum either
very wide, with 2 narrow medium teeth and > 8 lateral teeth of similar size (Figs. 5–6), or with 1 very broad median tooth and 5 laterals
(Fig. 4). Head dark brown, body ‘fleshy’ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diamesinae (Heptagyiini)
4.— Prementum with lamellae rather than brushes. Mentum usually narrower, with at most 3 median teeth. Body of various coloration . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Orthocladiinae
Notes: The only Tanypodinae recorded in the studied streams was Pentaneura sp. (Fig. 23). There was 1 specimen in the Surber samples of the
Glacier stream 14, site 3 (farthest from the glacier terminus), and it is apparently very rare in the region.
Subfamily Chironominae
1. Antennae long, placed on pedestals (Fig. 3) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tanytarsini
1.— Antenna shorter, growing directly from head (Chironomini, Pseudochironomini) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Mentum with a single broad median tooth, ventromental plates slender, scarcely separated medially. Anal tubules elongate, manifold
exceeding the length of parapods (Fig. 2) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Manoa
2.— Mentum with paired median teeth, first laterals shorter than median and second lateral teeth, ventromental plates widely separated.
Anal tubules shorter than parapods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polypedilum
Notes: The only Tanytarsini collected was 1 specimen of Tanytarsus sp. (Fig. 3) in stream 14, site 3. One individual of Manoa sp. (Pseudochirono-
mini) was recorded in Glacier stream 15, site 3. Both taxa are particularly rare.
Table 1. Basic environmental variables of the studied streams. For details see Kuhn et al. (2011).
Site code 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17
Humboldt
Yacupampa
Micahuayco
Micahuayco
Jatunhuayco
Jatunhuayco
Stream G 15 G 15 Gl 15 G 14 G 14 G 14 G 12 G 12 G 12
Spring stream
Rio Antisana
Site S1 S2 S3 S1 S2 S3 S1 S2 S3 S1 S2 S1 S2
Altitude (masl) 4,835 4,521 4,335 4,789 4,535 4,196 4,728 4,496 4,225 4,051 4,000 3,925 3,960 3,958 4,070 4,058 4,105
UTM 817040 815887 814882 816356 815255 811885 816375 815013 812563 810143 807352 808047 809341 808958 810177 810142 809926
Abbreviations: UTM – Universal Transverse Mercator coordinate system; AFDM – ash-free dry mass; CV – coefficient of variation.
2018 — Florida Entomologist — Volume 101, No. 4
Table 2. Chironomid taxa recorded in the surveyed streams in the Antisana region, Ecuador, in Jan 2008. Numbers indicate summary abundance in 5 Surber samples; + indicate records in qualitative samples.
Site code 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17
Micahuayco
Micahuayco
Rio Antisana
Jatunhuayco
Jatunhuayco
Spring stream
Stream G 15 G 15 Gl 15 G 14 G 14 G 14 G 12 G 12 G 12
Site S1 S2 S3 S1 S2 S3 S1 S2 S3 S1 S2 S1 S2
Chironominae
Manoa sp. — — 1 — — — — — — — — — — — — — —
Polypedilum sp. — — — — — 3 — — — + + + + + — — —
Tanytarsus sp. — — — — — 1 — — — — — — — — — — —
Orthocladinae
Corynoneura lobata gr. — — — — — 3 — — — — — — — + — — —
Cricotopus sp. A — 9 + — — — + + + — — — — — — — —
Cricotopus sp. B — 55 98 6 28 32 + 37 18 — + + + + + + +
Cricotopus sp. C — 2 7 — 15 230 — 1 88 + + + + + + + +
Cricotopus sp. D — — — — — 16 — — — + — — — — — — —
Limnophyes sp. — 6 — 1 40 7 2 5 3 + — + + + — — +
Metriocnemus fuscipes gr. — — 1 — — — — — — — — — — — — — —
Metriocnemus eurynotus gr. — — + — — — — — — — — — — — — — —
Onconeura sp. — — — — — — — — — + — — — — — — —
Orthocladius (Euo.) sp. — — — — — — — — — — + + + + + + +
Parakiefferiella sp. — — 1 — + 126 — — 2 + — — — — — — —
Pseudosmittia sp. — + — — — — — — — — — — — — — — —
Smittia sp. — — 2 — 2 — — — — — — — — + — — —
Podonominae
Parochlus sp. A 16 24 20 13 177 37 49 46 3 + + — + — + + +
Parochlus sp. B + 1 5 1 2 2 14 — — — — — — — — — —
Podonomus sp. 28 4 1 71 2 18 20 3 1 + + — — + — — +
Tanypodinae
Pentaneura sp. — — — — — 1 — — — — + — — — — — —
667
668 2018 — Florida Entomologist — Volume 101, No. 4
Figs. 2–7. Manoa sp.: 2a. Labrum-epipharyngis. 2b. Antennae. 2c. Mentum, mandibles and ventromental plates. 2d. Posterior parapods and anal tubules, lateral
view. Tanytarsus sp.: 3a Mandible. 3b. Mentum and ventromental plates. 3c. Head capsule, ventral view. 3d. Labro-epipharyngeal region (arrow indicates pre-
mandibles with 3 teeth). Limaya sp.: 4a. Mandible. 4b. Mentum. 4c. Antenna. 4d. Labro-epipharyngeal region. Paraheptagyia sp. A: 5a. Head capsule, dorsal view.
5b. Mentum and mandibles. 5c. Head capsule, ventral view, arrow indicates occipital margin. Paraheptagyia sp. B: 6a. Mentum. 6b. Mandibles. 6c. Anal end of body,
lateral view. 6d. Head capsule, ventral view, arrow indicates occipital margin. Parochlus sp. A: 7a, e. Anal end with posterior parapods, procercus and anal setae,
lateral and dorsal view. 7b. Mentum and mandibles. 7c. Claws of posterior parapods (note the 2 types of claws indicated by arrow). 7d. Antenna.
Figs. 8–13. Parochlus sp. B: 8a to c. Anal end with posterior parapod, procercus and anal setae, lateral (a, b) and dorsal view (c). Podonomus sp.: 9a. Anterior
parapod with fan shaped arrangement of claws, lateral view. 9b, c. Anal end with posterior parapod, procercus, and anal setae, dorsolateral view. 9d. Larvae under
low magnification. Corynoneura lobata group: 10a. Antenna. 10b. Mentum, mandibles, and labro-epipharyngeal region (arrows indicate bifid SI and pecten). 10c.
Head capsule, dorsolateral view. Cricotopus sp. A: 11a. Labro-epipharyngeal region. 11b. Body. 11c. Body setation (indicated by arrow). 11d. Head capsule, ventral
view (position of seta submenti indicated by arrow). 11e. Mandible. Cricotopus sp. B: 12a. Mandible. 12b. Head capsule, ventral view. 12c. Mentum view (position
of seta submenti indicated by arrow). 12d. Body segment with seta (indicated by arrow). Cricotopus sp. C: 13a. Mandible. 13b. Head capsule, ventral view. 13c.
Mentum (arrow indicates position of seta submenti).
Fig. 14–19. Cricotopus sp. D: 14a. Mentum. Arrow indicates the reduced second lateral tooth. 14b. Mandible. 14c. Head capsule, ventral view. Limnophyes sp.: 15a.
Anal end of body, anal tubules are indicated by arrow. 15b. Antenna and mandible, arrow indicates long antennal blade. 15c. Mentum with wedge shaped tubercle
(indicated by arrow). Metriocnemus eurynotus group: 16a. Mandible. 16b. Antenna. 16c. Mentum. 16d. Head capsule, ventral view. Metriocnemus fuscipes group:
17a. Antenna. 17b. Mandible. 17c. Mentum. Onconeura sp.: 18a. Head capsule. Arrow indicates antenna. 18b. Mentum and mandible. 18c. Antenna. 18d. Body
segments with strong setation. Orthocladius (Euo.) sp.: 19a. Mentum. 19b. Head capsule, ventral view. 19c. Detail of occipital margin. 19d. Antenna and madible.
Figs. 20-23. Parakiefferiella sp.: 20a. Mandible. 20b. Antenna. Arrow indicates vestigial sixth segment. 20c. Mentum. Pseudosmittia sp.: 21a. Head capsule, ventral
view. 21b. Anal end, ventral view. 21c. Mentum and mandible. Smittia sp.: 22a. Head capsule, ventral view. 22b. Mentum. 22c. Basally fused anterior parapods, with
2 types of claws (fine spines are indicated by arrow), lateral view. 22d. Posterior parapods (detail), ventral view. Arrow indicate a row of minute claws below regular
size claws. 22e. Anal end of the body, ventral view. Pentaneura sp.: 23a. Head capsule, ventral view. 23b. Ligula and paraligula. 23c. Maxillary palp with sensilla. 23d.
Anal end of body.
Notes: The most common orthoclads are Cricotopus sp. A, C. sp. B, and C. sp. C. Even though these morphotypes are very similar, they
can be distinguished as follows: Cricotopus sp. A (Fig. 11) has a characteristic body coloration; the first 3 body segments are pale, while
the remaining are purple. Seta submenti is situated between ventromental plates. The head is dark brown, the mentum, mandibles, and
occipital margin are dark brown to black. The apical tooth of mandible long and pointy. Abdominal segments with a pair of setal tufts (Fig.
11c) (also occurs in Cricotopus sp. B), but may be weak and easily overlooked. Cricotopus sp. A is common, most abundant in the uppermost
site, also occurs at lower elevations, but much less frequently. The most obvious difference between Cricotopus sp. B (Fig. 12) and C. sp. C
(Fig. 13) is the color and shape of occipital margin; although occipital margin of C sp. B is narrow and slightly darker than head capsule, C.
sp. C has broad and pale occipital margin, about the same color as head capsule. In addition, ventral opening of occipital margin in C. sp. C
has a characteristic trapezoidal shape, while that of C. sp. B is wide, without trapezoidal shape. Abdominal segments of C. sp. B bear very
weak setal tufts (Fig. 12d); however, they could be easily overlooked. Both types are common in the glacier-fed sites; however, C. sp. B is
most abundant in the middle ones, while C. sp. C is absent in the uppermost ones, and its abundance increases with decreasing altitude,
and is most abundant in the lowermost sites.
Subfamily Podonominae
1. Dark brown to black head capsule and procerci, claws of anterior and posterior parapods arranged in comb shape, anal setae compact
(Fig. 9). Common in all sites, most abundant in the uppermost ones, close to the glacier terminus . . . . . . . . . . . . . . . . . . . . . Podonomus
1.— Pale to light brown head capsule, procerci and body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Light brown body with obvious setae, procerci 3 to 4× as long as broad, anal setae arranged in fan shape, head slightly curved (Fig. 7) . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parochlus sp. A