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Corresponding author: Erika Mayumi Shimabukuro ( erika.msh@gmail.com ) Academic editor: Fabio Laurindo da Silva
© 2018 Erika Mayumi Shimabukuro, Susana Trivinho-Strixino.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Shimabukuro EM, Trivinho-Strixino S (2018) Madicolous Chironomidae from the Brazilian Atlantic Forest: a checklist with notes on altitudinal distributions (Diptera, Insecta). ZooKeys 751: 41-73. https://doi.org/10.3897/zookeys.751.20611
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Thin layers of water running over rocky surfaces are characteristic of madicolous habitats, which harbor a peculiar Chironomidae community. However, information on the identity, distribution, and ecology of madicolous chironomids in the Neotropical region are still sparse. The main purpose of this research is to reveal and contribute to the ecology of madicolous Chironomidae species, especially regarding their altitudinal distribution in the Atlantic Forest. Sampling was performed using our own designed emergence traps deployed from 0 to 2700 m a.s.l. in 70 sites in three mountains in southeastern Brazil. Sixty taxa of chironomids were collected and identified, of which only 22 are known to science. Most of the species showed a wider distribution than previously known, both in terms of geographic and altitudinal ranges, while others showed significant association with particular altitudinal bands (as evidenced by the indicator species analysis). Atlantic Forest mountainous regions are known to harbor one of the richest fauna in the world and have been suffering from several types of environmental impacts, including climate change, which will especially affect taxa living in specialized habitats. The narrow range of tolerance to environmental conditions verified for mountain species, and the fact that many of them are rare and endemic, make the conservation efforts in these areas indispensable.
hygropetric habitats, mountains, non-biting midges, semi-aquatic habitats, tropical forest
Madicolous habitats are characterized by a thin layer of water that frequently flows over rocky surfaces, and for this reason they are also known as hygropetric habitats. The first to use the term “hygropetrischen” was Thienemann in 1909, when studying the biology of trichopterans from Central Europe. Throughout the twentieth century, some catalogues of madicolous fauna were done in North America (
In natural ecosystems, madicolous insects can live in a wide range of habitats, such as shoreline of streams or in isolated overflowing groundwater. Additionally, when robust water bodies like streams and lakes are scarce, for example on mountaintops, madicolous biotopes can be the only source of permanent water allowing aquatic and semi-aquatic insects to establish themselves and survive, contributing to the maintenance of biodiversity in natural systems. The true madicolous inhabitants (eumadicoles) present morphological and physiological adaptations favoring their survival in such a specific environmental condition, as seen by the presence of strong locomotor appendages to hang on the rocky substrate in larval stages (
The Chironomidae family is one of the most diverse within Diptera. Species numbers reach an estimated 20,000 (
A high diversity of chironomids is expected to occur in natural madicolous habitats from tropical regions, but this biotope has so far been neglected in freshwater researches, making it difficult to have an estimate on the diversity of insects living in such habitats. Furthermore, concerning the taxonomy of chironomids, most of the descriptions are based exclusively on adult males, making it difficult to obtain the information on the habitats, behavior, and other ecological information related to aquatic stages.
Despite significant progress on Chironomidae research in the last decade (
The exceptional biodiversity verified in the Brazilian Atlantic Forest is mainly due to the environmental heterogeneity owed to its singular geographical characteristics. Specifically, the region is characterized by a high variation in latitudinal (originally from parallels 3° to 30°) and altitudinal ranges (0 to 2892 m a.s.l.). These generate a diverse forest composition and a wide variety of habitats (
This study was conducted inside three conservation units from the Atlantic Forest in southeastern Brazil: Serra do Mar State Park (PESM), Serra dos Órgãos National Park (PARNASO), and Serra da Mantiqueira Environmental Protection Area (APASM) (mean distance between those areas is 185 km) (Figure
PESM is the widest continuous protected area inside the Atlantic Forest (332,000 ha). It encompasses the whole coast of São Paulo State, including stretches at the sea level and some peaks that elevate the altitudinal range up to 1270 m a.s.l. The vegetation present in the region includes: mangroves, “restinga” (costal dunes), costal vegetation, ombrophilous dense forest, and “campos de altitude” (high altitude-vegetation composed predominantly of grass, shrub, and herbaceous vegetation). In this locality, sites between 0–1100 m a.s.l. were explored (Figure
PARNASO is in the mountainous region of Rio de Janeiro State, where it occupies 20,024 ha. The region’s relief is marked with slopes, which generates a high altitudinal gradient. The upmost site in the park reaches 2263 m a.s.l., and the vegetation changes alongside variations in elevation: submontane forest, montane forest, misty forest, and campos de altitude. In this park, intermediate altitudes were explored, between 1200–2100 m a.s.l. (Figure
APASM includes three states from the Southeast: São Paulo, Rio de Janeiro, and Minas Gerais, comprising an area of 421,804 ha. It harbors two of the five highest mountains in Brazil, including the culminant site at 2798 m a.s.l. (Pedra da Mina Mountain). The vegetation in the region creates a mosaic of phytophysiognomies, composed by upper-montane forests, araucaria forests and campos de altitude. In this locality, sites from 1700 to 2700 m a.s.l. were sampled (Figure
Madicolous habitats were sampled every 200–300 m along the gradient. At least three replicates were obtained in each altitudinal band.
The adult sampling was performed with emergence traps (
For each species found, a set of data from the literature was added, including previous information on their geographical distribution, altitudinal records, habitats and development stages and sexes so far known. After the literature data, the “Remarks” include the data obtained by us: 1. type of madicolous habitat where the species was found, such as stream edge (Figure
Information on the development stages, habitat, and locality of each species and morphospecies recorded in our study is summarized in Table 1 (supplementary material). When the development stage consists only of adults (A), it does not mean that immature specimens were not collected; rather, that the association between adult and immature stages were not yet established. In addition, some taxa were recorded only from its immature instars (I).
In order to test the degree of each species'affinity within the altitude in which they occurred, an indicator species analysis (
A total of 60 species, including 22 known species and 38 morphospecies, was recorded, as follows:
Two valid species; Neotropical; running water.
BRAZIL – Camacan, Bahia State (15°23'28"S, 39°33'56"W; 15°23'02"S, 39°34'10"W; 15°23'10"S, 39°34'03"W) Rio de Janeiro State (11°53'40"S, 45°36'06"W), Salesópolis, São Paulo State; Campos do Jordão, São Paulo State (22°41'40"S, 45°27'36"W); Grão-Pará, Santa Catarina State (28°11'28"S, 49°23'30"W); Urubici, Santa Catarina State (28°01'41"S, 49°22'36"W).
508–1014 m a.s.l.
Low-order streams.
L, P, M.
C. froehlichi was found on marginal rocks of small streams, at 206 m a.s.l. (23°27'52"S, 45°11'55"W) and 1444 m a.s.l. (22°26'51"S, 43°00'48"W), extending the altitudinal range of the species. Environmental characterization: Water temperature 10–17 °C; dissolved oxygen 9.0–9.2 mg.l-1; pH 5.0–5.5; dense canopy cover. The species was found in PESM (São Paulo State) and PARNASO (Rio de Janeiro State). Low altitudinal specificity (IndVal: 0.17; p = 0.84).
One valid species; Holartic, Nearctic, and Neotropical; standing water.
Unknown species. Lauterborniella sp. 1. Locality: PESM. Altitudinal record: 1080 m a.s.l. Significant altitudinal specificity (IndVal: 0.5; p = 0.02).
52 valid species; worldwide; running and stagnant water.
Unknown species. Nilothauma sp. 1 Locality: APASM. Altitudinal record: 1570 m a.s.l. Low altitudinal specificity (IndVal: 0.14; p = 1.00).
24 valid species; Neotropical, associated with freshwater sponges and submerged wood.
BRAZIL – Paru do Oeste River, missão Cururu, Amazonas State; Parque Estadual de Campos do Jordão, São Paulo State (22°41'40"S, 45°27'36"W).
20–1600 m a.s.l.
Larvae found in submerged wood in a rocky first-order stream.
L, P, M.
O. sublettei was found on marginal rocks of small streams, at 745 m a.s.l. Environmental characterization: Water temperature 17 °C; dissolved oxygen 8.2 mg.l-1; pH 5.5; fast flowing; dense vegetal canopy (more than 80% covered). The species was found in PESM (São Paulo State). Median altitudinal specificity (IndVal: 0.33; p = 0.14).
More than 440 valid species; worldwide; standing and running water.
BRAZIL – Tarumã River, Amazonas State; Florianópolis, Santa Catarina State (27°28'05"S, 48°22'58"W), UCAD, Santa Catarina State (27°31'51"S, 48°30'44"W), Santinho Beach, Santa Catarina State (27°27'S, 48°23'W).
20–80 m a.s.l.
Adults collected close to a large Amazonian river. Larva found in leaf packs and detritus associated with the following bromeliad species: Aechmea lindeni (E. Morren) Baker, Canistrum lindenii (Regel) Mez, Neoregelia laevis (Mez) L.B. Smith, Nidularium innocentii Lem., Vriesea philippocoburgii Wawra, and V. vagans (L.B. Smith) L.B. Smith.
L, P, F, M.
P. solimoes was found on marginal rocks of small stream, at 1570 m a.s.l., extending the elevation records of this species. Environmental characterization: Water temperature 16.3 °C; dissolved oxygen 8.4 mg.l-1; pH 6.4; slow flowing; vegetal canopy absent. The species was found in APASM (Minas Gerais State). Low altitudinal specificity (IndVal: 0.14; p = 1.0).
Unknown species. Polypedilum (s. str.) sp. 1. Locality: APASM. Altitudinal record: 1570 m a.s.l. Low altitudinal specificity (IndVal: 0.14; p = 1.00); Polypedilum (s. str.) sp. 2 Locality: PESM. Altitudinal record: 1080 m a.s.l. High altitudinal specificity (IndVal: 0.33; p = 0.06); Polypedilum (s. str.) sp. 3. Locality: PARNASO. Altitudinal record: 2125 m a.s.l. Low altitudinal specificity (IndVal: 0.17; p = 0.84); Polypedilum (Pentapedilum) sp. 1; Locality: PESM. Altitudinal record: 200 m a.s.l. Median altitudinal specificity (IndVal: 0.33; p = 0.15). Polypedilum (Pentapedilum) sp. 2. Locality: PARNASO. Altitudinal record: 2125 m a.s.l. Low altitudinal specificity (IndVal: 0.17; 6 p = 0.85). Polypedilum (Tripodura) sp. 1. Locality: PESM. Altitudinal record: 25 m a.s.l. Low altitudinal specificity (IndVal: 0.25; p = 0.33).
101 valid species; worldwide; miners of living or dead vegetal tissue.
Unknown species. Stenochironomus sp. 1. Locality: PESM. Altitudinal record: 1075 m a.s.l. Low altitudinal specificity (IndVal: 0.17; p = 0.84); Stenochironomus sp. 2. Locality: PESM. Altitudinal record: 70 m a.s.l. Low altitudinal specificity (IndVal: 0.25; p = 0.32).
13 valid species; Nearctic, Palearctic, Neotropical; running and standing water.
P. ruah was recorded for the first time on marginal rocks of a small stream, at 2575 m a.s.l. (22°25'41"S, 44°50'38"W), in APASM (Minas Gerais State). Larva, pupa, female and male have been described by
45 valid species; worldwide; running and stagnant water
BRAZIL – Parque Estadual do Jaraguá, São Paulo State (23°24'S, 45°44'W).
800 m a.s.l.
Stream.
M.
P. silentii was found on marginal rocks of small streams, from 200–1445 m a.s.l., extending the altitudinal range of the species. Environmental characterization: Water temperature varied from 10–21 °C; dissolved oxygen 7.9–9.9 mg.l-1; pH 5–5.5; slow to fast flowing; occurring in scarce vegetal canopy to dense coverage (30–70% covered). The species was found in PARNASO (Rio de Janeiro State) and PESM (São Paulo State). Low altitudinal specificity (IndVal: 0.24; p = 0.25).
More than 300 valid species; worldwide; aquatic and terrestrial
T. alaidae was recorded for the first time on marginal rocks of a small stream, at 2575 m a.s.l. (22°25'41"S, 44°50'38"W), in APASM (Minas Gerais State). At present, only adult males have been described (
T. alienus was recorded for the first time on marginal rocks of a small stream, at 2575 m a.s.l. (22°25'41"S, 44°50'38"W), in APASM (Minas Gerais State). At present, only adult males have been described (
T. angelae male adults were recorded in a temporary pool at 2218 m a.s.l. (22°24'697"S, 44°50'93"W) in APASM (Minas Gerais State), in a rocky fountain at 2126 m a.s.l. (22°27'60.5'S, 43°01'68.9"W) in PARNASO (Rio de Janeiro State), and in the same locality at 1444 (rocky substrate marginal to a stream – 22°26'84.8'S, 43°00'79.8"W). At present, only adult males have been described (
BRAZIL – Paquequer River, Teresópolis, Rio de Janeiro State.
1100 m a.s.l.
Adults collected in light trap close to a first-order stream.
M.
T. digitatus was found on marginal rocks of small streams, at 25 m (23°28'20.72"S, 45°11'38.43"W) and 1445 m a.s.l (22°26'84.8"S, 43°00'79.8"W), extending the altitudinal range of the species. Environmental characterization: Water temperature 10–19.5 °C; dissolved oxygen 8.1–9.0 mg.l-1; pH 5.0; slow flowing; dense canopy cover. The species was found in PESM (São Paulo State) and PARNASO (Rio de Janeiro State). Low altitudinal specificity (IndVal: 0.5; p = 0.02).
BRAZIL – São Carlos, São Paulo State (21°59'10"S, 47°52'32"W; 21°58'16"S, 47°53'14"W; 21°59'21.4"S, 47°51'14.2"W); Corumbá, Mato grosso do Sul State (19°34'30.06"S, 57°00'52.4"W).
90–850 m a.s.l.
Low-order streams and eutrophic lake.
L, P, M.
T. giovannii was found on marginal rocks of small streams, at 2575 m a.s.l., extending the altitudinal range of the species. Environmental characterization: Water temperature 22 °C; dissolved oxygen 7.4 mg.l-1; pH 5.5; fast flowing; vegetal canopy absent. The species was found in APASM (Minas Gerais State). Low altitudinal specificity (IndVal: 0.17; p = 0.84).
Approximately 96 valid species; worldwide; running and stagnant waters.
BRAZIL – Parque Estadual do Jaraguá, São Paulo State (23°27'S, 46° 45'W) Serra do Japi, Jundiaí, São Paulo State (23°14'30"S, 46°57'16"W); Paraisópolis, Minas Gerais State (22°39'54.81"S, 45°55'38.29"W); São Luís do Purunã, Paraná State (25°27.180'S, 49°43.435'W); BRAZIL, Serra da Bodoquena, Mato Grosso do Sul State (20°41'49"S, 56°52'54"W); Alto Paraíso de Goiás, Goiás State (14°9'34.92"S, 47°35'37.32W); COSTA RICA, Caccao, Guanacaste.
750–1370 m a.s.l.
Litter in stones marginal to small streams.
L, P, F, M.
C. unicapsulata was found on marginal rocks of small streams, at 2575 m a.s.l., extending its altitudinal occurrence. Environmental characterization: Water temperature 22 °C; dissolved oxygen 7.4 mg.l-1; pH 5.5; slow flow to stagnant; vegetal canopy absent (shrub-herbaceous vegetation. The species was found in APASM (Minas Gerais State). Low altitudinal specificity (IndVal: 0.17; p = 0.85).
BRAZIL – Ubatuba, São Paulo State (23°30.468'S, 45°11.923'W and 23°30.789'S, 45°14.442'W)
0–60 m a.s.l.
Litter standing near the water surface from small streams.
L, P, F, M.
C. hermanni was found on marginal rocks of small streams and rocky seepages (large exposed rock in the middle of the forest), at 1570–1580 m a.s.l., extending the altitudinal occurrence of this species. Environmental characterization: Water temperature 15–16.3 °C; dissolved oxygen 8.4–10.2 mg.l-1; pH 5.0–6.4; slow to fast flowing; vegetal canopy completely absent. The species was found in APASM (Minas Gerais State) and PARNASO (Rio de Janeiro State), extending slightly the geographical distribution of this species. Low altitudinal specificity (IndVal: 0.29; p = 0.23).
BRAZIL – Parque Estadual do Jaraguá, São Paulo State (23°27'S, 46°45'W), Nova Friburgo, Rio de janeiro State, Rio Boa Vista; Bocaina de Minas, Minas Gerais State (22°19'S, 44°34'W); Serra do Japi, Jundiaí, São Paulo State (23°14'30"S, 46°57'16"W); Serra da Bodoquena, Mato Grosso do Sul State (20°41'49"S, 56°52'54"W).
700–1200 m a.s.l.
Litter near the water surface of a small shallow stream.
L, P, F, M.
C. septadentata was found on marginal rocks of small streams and on rocky seepages (large exposed rock in the middle of the forest), from 1045–1580 m a.s.l., extending slightly the altitudinal range of this species. Environmental characterization: Water temperature 15–17 °C; dissolved oxygen 8.6–10.2 mg.l-1; pH 5; moderate to fast flowing; vegetal canopy reduced or absent (less than 30% covered). The species was found in PARNASO (Rio de Janeiro State) and PESM (São Paulo State). Low altitudinal specificity (IndVal: 0.09; p = 0.88).
BRAZIL – PESN, Ubatuba, São Paulo State (23°30.789'S, 45°14.442'W; 23°31.068'S, 45°14.845'W; 23°31.231'S, 45°14.625'W); Alto Paraíso de Goiás, Goiás State (14°9'34.92"S, 47°35'37.32"W); Serra da Bodoquena, Mato Grosso do Sul State (20°41'49"S, 56°52'54"W), São Simão, São Paulo State; São Luís do Purunha, Paraná State (25°27.180'S, 49°43.435'W).
20–1020 m a.s.l.
Surface of stones of shallow fast flowing waters and also in litter near the water surface of streams.
L, P, F, M.
C. sertaodaquina was found on marginal rocks of small streams, at 70 m a.s.l. Environmental characterization: Water temperature 17 °C; dissolved oxygen 9.2 mg.l-1; pH 5.5; wet substrate, without any flow; reduced vegetal canopy (about 20% covered). The species was found in PESM (São Paulo State). Low altitudinal specificity (IndVal: 0.25; p = 0.32).
Seven valid species; Neotropical and Nearctic; running and stagnant water.
BRAZIL, Serra do Japi, Jundiaí, São Paulo State (23°14'38"S, 46°57'02"W); PESM, Ubatuba, São Paulo State (23°30.46'S, 45°11.923'W and 23°30.789'S, 45°14.442'W).
1058 m a.s.l.
Litter below a waterfall of a first-order stream.
L, P, F, M.
O. japi was found on marginal rocks of small streams, at 1570 m a.s.l., extending the altitudinal record of the species. Environmental characterization: Water temperature 16.3 °C; dissolved oxygen 8.4 mg.l-1; pH 6.4; slow flowing; vegetal canopy completely absent. The species was found in APASM (Minas Gerais State), extending the geographical distribution of the species. Low altitudinal specificity (IndVal: 0.14; p = 1.0).
BRAZIL – PESM, Ubatuba, São Paulo State (23°31.068'S, 45°14.845'W; 23°31.087'S, 45°14.621'W; 23°30.789'S, 45°14.442'W); São Francisco de Paula, Rio Grande do Sul State.
23–61 m a.s.l.
Surface of stones in fast flowing waters.
L, P, F, M.
O. oncovolsella was found on marginal rocks of small streams, at 1445 m a.s.l., extending the altitudinal record of the species. Environmental characterization: Water temperature 10 °C; dissolved oxygen 9 mg.l-1; pH 5; fast flowing; partial vegetal canopy (50% covered). The species was found in PARNASO (Rio de Janeiro State). Low altitudinal specificity (IndVal: 0.25; p = 0.31).
Unknown species. Onconeura sp. 1. Locality: PARNASO. Altitudinal record: 1445 m a.s.l. Low altitudinal specificity (IndVal: 0.25; p = 0.30); Onconeura sp. 2 Locality: PARNASO. Altitudinal record: 1445 m a.s.l. Low altitudinal specificity (IndVal: 0.25; p = 0.32); Onconeura sp. 3 Locality: PESM. Altitudinal record: 1085 m a.s.l. Low altitudinal specificity (IndVal: 0.17; p = 0.83).
53 valid species; worldwide; running and stagnant water.
Unknown species. Thienemanniella sp.1. Locality: APASM. Altitudinal record: 1570 m a.s.l. Low altitudinal specificity (IndVal: 0.14; p = 1.00).
115 valid species; worldwide; terrestrial and semi-terrestrial, few aquatic.
BRAZIL – Parque Estadual Intervales, Iporanga, São Paulo State (24°30'S, 48°37'W); PANAMA, Canal Zone, Curundu, Holbrook Air Force Base.
20–100 m a.s.l.
Mosses on tree barks.
L, P, F, M.
B. carus was found on marginal rocks of small streams, at 1075 m a.s.l., extending the altitudinal range of the species. Environmental characterization: Water temperature 15.8 °C; dissolved oxygen 8.1 mg.l-1; pH 5; moderate to fast flowing; dense vegetal canopy (more than 70% covered). The species was found in PESM (São Paulo State). Low altitudinal specificity (IndVal: 0.17; p = 0.85).
Unknown species. Bryophaenocladius sp. 1. Locality: PESM. Altitudinal record: 1075 m a.s.l. Low altitudinal specificity (IndVal: 0.17; p = 0.87).
Four valid species; Brazil, Atlantic Forest; unknown habitat, but possibly terrestrial or semi-terrestrial.
BRAZIL – Parque Estadual Intervales, Ribeirão Grande, São Paulo State (24°15'S, 48°10'W).
500 m a.s.l.
Unknown, but possibly terrestrial or semi-terrestrial.
M.
C. intervales was found on marginal rocks of small streams, at 740 m a.s.l., extending slightly the altitudinal range of this species. Environmental characterization: Water temperature 15.7 °C; dissolved oxygen 9.4 mg.l-1; pH 5.5; very slow flowing; partial vegetal canopy (50% covered). The species was found in PESM (São Paulo State). Median altitudinal specificity (IndVal: 0.33; p = 0.14).
218 valid species; worldwide; running and standing water.
Unknown species. Cricotopus sp. 1. Locality: APASM. Altitudinal record: 2575 m a.s.l.. Low altitudinal specificity (IndVal: 0.17; p = 0.85); Cricotopus sp. 2 Locality: APASM and PARNASO. Altitudinal range: 1445–1570 m a.s.l. Low altitudinal specificity (IndVal: 0.15; p = 0.47); Cricotopus sp. 3 Locality: PESM. Altitudinal range: 70–1075 m a.s.l. Low altitudinal specificity (IndVal: 0.13; p = 0.58); Cricotopus sp. 4 Locality: PESM. Altitudinal range: 70–200 m a.s.l.. Significant altitudinal specificity (IndVal: 0.71; p = 0.003); Cricotopus sp. 5 Locality: APASM. Altitudinal range: 1750 m a.s.l. Low altitudinal specificity (IndVal: 0.14; p = 1.0).
91 valid species; worldwide, except for Oceania and Antarctic; aquatic, terrestrial and semi-terrestrial habitats.
BRAZIL – Serra do Corvo Branco, Grão-Pará, Santa Catarina State (28°03'21"S, 49°22'00"W).
1241 m a.s.l.
Madicolous habitat.
L, P, F, M.
L. guarani was found on rocky seepages and also on marginal rocks of first order streams, from 1570–2700 m a.s.l., extending the altitudinal range of the species. Environmental characterization: Water temperature 16.3–22 °C; dissolved oxygen 6.4–8.4 mg.l-1; pH 5.5–6.4; slow to fast flowing; vegetal coverage completely absent (shrub-herbaceous vegetation). The species was found in APASM (Minas Gerais State), extending the geographical records to northernmost. Median altitudinal specificity (IndVal: 0.19; p = 0.18).
BRAZIL – Parque João Paulo II, Curitiba, Paraná State; UCAD, Florianópolis, Santa Catarina State; Parque Nacional de São Joaquim, Urubici, Santa Catarina State (28°07'32"S, 49°29'38"W); Nova Teutônia, Santa Catarina State (27°11'S, 52°23'W).
300–1822 m a.s.l.
Adults collected with entomological net and malaise trap close to stream.
F, M.
L. gercinoi was found on marginal rocks of small streams, from 1080–1445 m a.s.l. Environmental characterization: Water temperature 10.0–16.1 °C; dissolved oxygen 8.1–9.9 mg.l-1; pH 5; slow to fast flowing; partial canopy (about 50–70% covered). The species was found in PARNASO (Rio de Janeiro State), PESM (São Paulo State) extending the geographical records to northernmost. Low altitudinal specificity (IndVal: 0.19; p = 0.2).
Unknown species. Limnophyes sp. 1. Locality: PARNASO. Altitudinal range: 1445–2125 m a.s.l. Low altitudinal specificity (IndVal: 0.11; p = 0.87).
Four valid species; Nearctic and Neotropical; unknown habitat, probably semi-aquatic and terrestrial.
BRAZIL – Parque Nacional de São Joaquim, Urubici, Santa Catarina State (28°07'37"S, 49°28'47"W).
1670 m a.s.l.
Male collected in malaise trap in cloud forest.
M.
L. biancae was found on marginal rocks of low order streams and small waterfalls, extending the altitudinal records from 1570–2575 m a.s.l. Environmental characterization: Water temperature varied from 11–22 °C; dissolved oxygen 7.4–10.0 mg.l-1; pH 4.5–6.4; slow to fast flowing; vegetal coverage completely absent (shrub-herbaceous vegetation). The species was found in PARNASO (Rio de Janeiro State) and APASM (Minas Gerais State), the northernmost records. Low altitudinal specificity (IndVal: 0.16; p = 0.39).
Unknown species. Lipurometriocnemus sp. Locality: APASM. Altitudinal record: 2700 m a.s.l. Low altitudinal specificity (IndVal: 0.2; p = 0.45).
67 valid species, worldwide, except for Oceania and Antarctic; mosses, Phytotelmata, springs, ditches, streams, lakes, and rock pools.
Unknown species. Metriocnemus sp. 1 Locality: APASM. Altitudinal record: 2200 m a.s.l. Low altitudinal specificity (IndVal: 0.17; p = 0.84).
44 valid species; worldwide; running and standing waters.
BRAZIL – Taquara, Rio Grande do Sul State (29°46'S, 50°53'W); São Francisco de Paula, Rio Grande do Sul State (29°26'S, 50°35'W); Bom Jesus, Rio Grande do Sul State (28°40'S, 50°26'W).
600–1000 m a.s.l.
Stream.
P, M.
P. strixinorum was found on marginal rocks of small streams, at 1045 m a.s.l., extending slightly the altitudinal range of the species. Environmental characterization: Water temperature 17 °C; dissolved oxygen 8.6 mg.l-1; pH 5; fast flowing; reduced vegetal canopy (less than 20% covered). The species was found in PESM (São Paulo State) extending the geographical records to northernmost. Low altitudinal specificity (IndVal: 0.17; p = 0.85).
Unknown species. Parakiefferiella sp. 1. Locality: PESM. Altitudinal record: 70 m a.s.l. Low altitudinal specificity (IndVal: 0.25; p = 0.33).
34 valid species; worldwide; springs, streams and rivers.
Unknown species. Parametriocnemus sp. 1. Locality: APASM. Altitudinal record: 2575 m a.s.l. High altitudinal specificity (IndVal: 0.33; p = 0.06); Parametriocnemus sp. 2. Locality: PESM and APASM. Altitudinal range: 25–1570 m a.s.l. Low altitudinal specificity (IndVal: 0.12; p = 0.70); Parametriocnemus sp. 3. Locality: PESM and APASM. Altitudinal range: 25–1445 m a.s.l. Low altitudinal specificity (IndVal: 0.11; p = 0.76).
98 valid species; worldwide; aquatic, terrestrial and semi-terrestrial habitats.
BRAZIL – Parque Nacional de São Joaquim, Urubici, Santa Catarina State (28°07'32"S, 49°29'38"W).
1822 m a.s.l.
Male collected in malaise trap in cloud forest, close to small stream.
M.
P. catarinense was found on rocky seepages, at 2200 m a.s.l., extending slightly it altitudinal occurrence. Environmental characterization: water temperature 27.6 °C; dissolved oxygen 7.0 mg.l-1; pH 6; slow flowing; vegetal coverage completely absent (shrub-herbaceous vegetation). The species was found in APASM (Minas Gerais State), extending the geographical records to northernmost. Low altitudinal specificity (IndVal: 0.17; p = 0.84).
69 valid species; worldwide except Antarctica and Oceania; mostly rheophilic.
Unknown species. Rheoricotopus sp. 1 Locality: APASM. Altitudinal record: 2200 m. a.s.l. Low altitudinal specificity (IndVal: 0.17; p = 0.85); Rheoricotopus sp. 2. Locality: PARNASO. Altitudinal range: 1580–1670 m a.s.l. Median altitudinal specificity (IndVal: 0.29; p = 0.11).
One valid species; Brazil, Atlantic Forest; unknown habitats.
BRAZIL – Parque Nacional de São Joaquim, Urubici, Santa Catarina State (28°07'37"S, 49°28'47"W).
1670 m a.s.l.
Male collected in malaise trap in cloud forest.
M.
U. montana was found on rocky seepages, from 2200–2700 m a.s.l., expanding its altitudinal range. Environmental characterization: Water temperature varied from 21.3–27.6 °C; dissolved oxygen 6.4–7.0 mg.l-1; pH 6; slow flowing; vegetal coverage completely absent (shrub-herbaceous vegetation). The species was found in APASM (Minas Gerais State), extending the geographical records to northernmost. Median altitudinal specificity (IndVal: 0.26; p = 0.13).
Unknown species. Urubicimbera sp. 1. Locality: APASM. Altitudinal range: 2575–2700 m a.s.l. Significant altitudinal specificity (IndVal: 0.91; p = 0.002).
40 valid species; Neotropical and Australasian; running water and tarn inhabitants.
P. mina was recorded for the first time on marginal bedrock of a mountain stream, at 1270 m a.s.l. (20°25'12"S, 41°50'45.6"W), in Serra do Caparaó (Espírito Santo State) (
BRAZIL – Mantiqueira and Espinhaço mountain ranges: Monte Verde, Minas Gerais State (22°53'9.6"S, 46°1'55.2"W); Campos do Jordão, São Paulo State (22°46'1.2"S, 45°31'15.6"W); Teresópolis, Rio de Janeiro State (22°27'3.6"S, 43°0'50.4" W); Alto Caparaó, Minas Gerais State (20°25'12"S, 41°50'45.6"W).
1275–1815 m a.s.l.
Pupae found in a first-order stream; larvae found in madicolous habitats.
L, P, M, F.
In this study larvae was found living on marginal rocks of a low order stream and in isolated rocky seepages, extending the altitudinal records up to 2700 m a.s.l. Environmental characterization: Water temperature varied from 10–22 °C; dissolved oxygen 6.4–9.0 mg.l-1; pH 5.0–6.0; very slow water flow; vegetal coverage completely absent (shrub-herbaceous vegetation). P. pepinellii was found in PARNASO (Rio de Janeiro State) and APASM (Minas Gerais State). Low altitudinal specificity (IndVal: 0.25; p = 0.32).
Four valid species; Nearctic and Neotropical; madicolous.
BRAZIL, Ubatuba, São Paulo State, (23°30.468'S, 45°11.923'W)
0 m a.s.l.
Few larvae found on leaf litter in shallow-water streams flowing over granite outcrops.
L, P, M.
H. caissara was found on marginal rocks of a small stream, at 200 m a.s.l. This slightly extended the altitudinal records of the species. Environmental characterization: Water temperature 21 °C; dissolved oxygen 7.9 mg.l-1; pH 5.5; fast flowing; sparse vegetal canopy (less than 30% covered). The species was found in PESM (São Paulo State). Median altitudinal specificity (IndVal: 0.33; p = 0.13).
Unknown species. Hudsonimyia sp.1. Locality: PESM. Altitudinal record: 1080 m a.s.l. Low altitudinal specificity (IndVal: 0.17; p = 0.85).
A summarized list of the species, morphospecies and the genera of immature found, along with respective ecological and geographical information, is presented in supplementary material (Table 1). In this study, the chironomid community was predominantly composed of species belonging to the subfamily Orthocladiinae (35 spp.), followed by Chironominae (21 spp), Podonominae and Tanypodinae (2 spp each). Among the 60 species recorded, a higher percentage has been found at APASM (45%), of which 85% were exclusive from this locality. Further, 31% of the possible new species occurred above 2100 m a.s.l. Only five from the 60 species recorded were significant indicators of specific altitudes, they are: Urubicimbera sp. 1, Cricotopus sp. 4, Pseudochironomus ruah, Lauterborniella sp. 1 and Podonomus mina (Figure
Regarding the 22 recognized species, 17 of them had spread the altitudinal distribution (Figure
For those species that have so far been verified at low altitudes, such as Tanytarsus giovannii, Polypedilum solimoes, Onconeura oncovolsella, Corynoneura hermanni and Bryophaenocladius carus, the amplitude of the altitudinal distribution was even more remarkable, adding more than 1500 m to the altitudinal range in some cases. The only exception was Hudsonimyia caissara, that have been firstly reported at the sea level, and here it was found at 200 m a.s.l., slightly increasing the altitudinal range of the species (Figure
Compositional changing in chironomid assemblages along altitudinal gradients have been verified by many researchers worldwide (
A clear gap on the taxonomic knowledge of mountain fauna can be observed. This gap is likely due to the low accessibility of these areas, thereby hampering sampling strategies. Studies in mountain regions are urgently needed, especially when dealing with one of the most threatened biomes in the world (
The indicator's analysis evidenced that all species significantly associated with their respective altitudinal band were previously unknown to science. All of them, except for Cricotopus sp. 4, recorded from 70–200 m a.s.l. were found exclusively at mountaintops. Lauterborniella sp. 1 was recorded at the highest sites in PESM and the remaining taxa were recorded at the summit of APASM mountains. Regarding the geophysical characteristics of mountaintops (shape, size, insulation value), also known as Inselbergs (
Our new records extend the altitudinal range of 17 known species. Most species seem to tolerate a wide altitudinal range, such as Tanytarsus giovannii and Limnophyes gercinoi, while others presented a narrow range, such as Hudsonimyia caissara. The altitudinal range is related to the extent of the geographical distribution of each species; species that are widely distributed are expected to occur in a wider range of altitudes compared to those that have limited distribution (
Madicolous habitats have never been formally studied in Brazilian mountainous regions, in contrast to other Atlantic Forest water bodies in which the Chironomidae fauna have already been extensively investigated (
Despite the low knowledge on semi-aquatic forms, evidences from chironomids fossils preserved in amber reveals that terrestrial life-styles have been common since the late Eocene (about 40 million years ago) (
Only two of the species verified here were previously known to occur in madicolous habitats (Podonomus pepinellii and Limnophyes guarani). Although Podonomus larvae can be found in streams and other fast flowing running waters, they are also common on the edge of streams (
The larvae of Hudsonimyia caissara were originally found in low abundance (two specimens) in leaf litter of a mountain stream (
It is not surprising that members of Hudsonimyia, Bryophaenocladius, Metriocnemus, Limnophyes and Pseudosmittia have been found in madicolous habitats during this study. These genera are known to have larval instars associated with semi-aquatic and terrestrial conditions.
Although many genera were expected to occur, some were particularly intriguing, such as Stenochironomus and Oukuriella. Both are known to be highly habitat-specialized in larval phase. The first is a vegetal miner (
Far from being semi-aquatic, most of the Rheotanytarsus species require flowing water conditions to survive and emerge (Coffman and Ferrington 1996). However, the capacity to live in madicolous habitats may not be disregarded as some species, such as R. gloveri, demonstrated tolerance to drying rock faces of streams and survived in thin layers of current water (
The procedure of rearing immature specimens to obtain the adults is most of the time unsuccessful due to their environmental requirements (
Using our modified emergence traps allowed us to assure that the immature organisms and the adults from the species sampled in this study were madicolous inhabitants. The association and description of the immature is a fundamental task when studying chironomids, best accomplished with the help of molecular tools, such as DNA barcode. The larva and the pupa of P. silentii have been successfully associated with adult males after this investigation (
We thank CNPq for financial support: PhD scholarship and productivity grant (141031/2013–1 and 306402/2010–6, respectively). We also thank Dr. Luiz Carlos de Pinho, Dr. Sofia Wiedenbrug, and Dr. Livia Fusari for taxonomic assistance, and Gilmar Perbiche Neves and Victor Saito for their help with field work.
Table S1. Taxa recorded in madicolous habitats from Atlantic Forest in Southeastern Brazil