Research Article
Print
Research Article
Poxyaibamberus Andersen & Dantas, gen. nov. (Diptera, Chironomidae, Orthocladiinae) from Brazil
expand article infoTrond Andersen, Galileu P. S. Dantas§, Viktor Baranov|, Annui M. Sanz-laParra, Humberto F. Mendes, Neusa Hamada§
‡ University of Bergen, Bergen, Norway
§ Instituto Nacional de Pesquisas da Amazônia, Coordenação de Biodiversidade (CoBio), Manaus, Brazil
| Estación Biológica de Doñana-CSIC / Doñana Biological Station-CSIC, Seville, Spain
¶ Universidade Federal de Alfenas, Alfenas, Brazil
Open Access

Abstract

Poxyaibamberus Andersen & Dantas, gen. nov. is erected based on the males of two species, P. jamanximensis Andersen & Dantas, sp. nov. from Jamanxim National Park, Pará State, Brazil, and P. ubajarensis Andersen & Dantas, sp. nov. from Ubajara National Park, Ceará State, Brazil. Both species have a comparatively short and wide head, with large eyes and short, five-segmented palps; a strong subapical seta on the ultimate flagellomere; scalpellate acrostichals; no setae on the wing veins except for one seta on the brachiolum; a long costal extension; and a large triangular anal point and a very long heel on the gonostylus. The systematic position of the new genus is briefly discussed.

Key words

Aquatic insects, new species, Neotropical Region, taxonomy

Introduction

The number of orthoclad species known from Brazil has increased rapidly during the last three decades. In their catalog of the Neotropical and Mexican chironomids Spies and Reiss (1996) recorded eight Orthocladiinae species in six genera from Brazil; one of these, Ichthyocladius neotropicus Fittkau, 1974, was listed as uncertain and has later been proven to not occur in Brazil (Mendes et al. 2004). Today more than 144 species in 44 genera are known to occur in the country (Pinho et al. 2024), although there is still a significant gap in the North and Northeast regions, where only 23 and 14 species of Orthocladiinae have been recorded. Many orthoclads encountered in Brazil do not readily fit into any described genus, and many new genera have been described. Several genera, namely Gynocladius Mendes, Sæther & Andrade-Morraye, 2005; Oleia Andersen & Mendes, 2007; Saetherocladius Andersen & Mendes, 2007; Saetherocryptus Andersen & Mendes, 2007; Saetherolabis Andersen & Mendes, 2007; Saetherops Andersen & Mendes, 2007; Lyrocladius Mendes & Andersen, 2008; Ubatubaneura Wiedenbrug & Trivinho-Strixino, 2009; Iporangomberus Mendes & Andersen, 2012; Pebapomberus Mendes & Andersen, 2012; Miambera Andersen & Mendes, 2012; Maximberus Andersen & Mendes, 2012; Jururumberus Mendes & Andersen, 2013; Uirassubrillia Mendes, Andersen & Pinho, 2013; Caaporangombera Andersen, Pinho & Mendes, 2015; Mariambera Andersen, Mendes & Pinho, 2015; and Urubicimbera Andersen, Pinho & Mendes, 2015 are so far endemic to Brazil (Mendes et al. 2005, 2013; Andersen and Mendes 2007, 2012a, 2012b; Mendes and Andersen 2008, 2012a, 2012b, 2013; Wiedenbrug and Trivinho-Strixino 2009; Andersen et al. 2015a, 2015b, 2015c). However, several of these genera are expected to be more widely distributed when the chironomid fauna of neighboring countries is better studied.

Below we describe a new genus based on the males of two new species collected in Pará and Ceará states in northern and northeastern Brazil. Both species have a comparatively wide head with large eyes and short, five-segmented palps, a strong subapical seta on the ultimate flagellomere, scalpellate acrostichals, a long costal extension, no setae on the wing membrane and veins except for one seta on the brachiolum, a large triangular anal point, and a gonostylus with a very long heel. The systematic position of the new genus is briefly discussed.

Materials and methods

The specimens were collected with Shannon traps (Shannon 1939) or light traps and preserved in 80% ethanol during the fieldwork. Prior to examination they were mounted in Euparal following the procedure outlined by Sæther (1969). Morphological terminology follows Sæther (1980). Coloration is based on the slide mounted specimen.

For the phylogenetic analysis we have used a morphological character matrix with 45 taxa and 83 characters. Characters were sampled from larvae, pupae, and adult males and females. Fossils were of course underrepresented in terms of characters available for observation (see https://github.com/chironomus/Poxyaibamberus-). The character matrix for the phylogenetic analysis was built using NEXUS DATA EDITOR v. 0.5. 0 (Page 2001).

First, a Bayesian analysis of the morphological matrix alone in MRBAYES 3.2.2. (Ronquist et al. 2012) was conducted using the Bayesian implementation of Lewis’ Markov models (Lewis 2001). In Bayesian inference, two Markov chains were run simultaneously for 10 million generations using a discrete Dirichlet distribution with equal state frequencies (Lewis 2001). Substitution model was set to “gamma” (Nylander et al. 2004). The first 25 0000 generations were discarded as a burn-in (number of MRBAYES generations of the tree topology before the apparent stationary condition) (Nylander et al. 2004). Consensus trees showing all compatible groups and 50% compatible groups were computed in MRBAYES. Ancestral character state analysis was conducted on the allcompat consensus tree based on morphology alone using ANCTRESH in the PHYTOOLS package v. 0.7-80 (Revell 2012). R code for this analysis, alongside the data is provided in https://github.com/chironomus/Poxyaibamberus-.

To deal with the uncertainty of the positions of the genus on the tree caused by the lack of knowledge of character states, the Klopfstein and Spasojevic ROGUEPLOTS approach was applied (Klopfstein and Spasojevic 2019). A morphological allcompat tree from MRBAYES (as described above) was used to place Poxyaibamberus into the high posterior probability regions. ROGUEPLOTS for every species and accompanying R code are provided in the https://github.com/chironomus/Poxyaibamberus-.

Both holotypes are kept in the Invertebrate collection at the Instituto Nacional de Pesquisas da Amazônia (INPA), Manaus, Brazil.

Taxonomic account

Family Chironomidae Newman, 1834

Subfamily Orthocladiinae Kieffer, 1911

Poxyaibamberus Andersen & Dantas, gen. nov.

Type species

Poxyaibamberus jamanximensis Andersen & Dantas, sp. nov.

Diagnosis

Small species, wing length 1.1–1.3 mm.

Description

Male antenna with 13 flagellomeres, strongly plumose, groove beginning on flagellomere 4, few sensilla chaetica apparently only present on flagellomere 13, with strong subapical seta. Antennal ratio 0.9.

Head short and wide. Eye bare, large, reniform, without dorsomedian extension. Temporal setae in single row, consisting of inner and outer verticals. Frontal tubercle absent. Tentorium and stipes normal. Clypeus with few setae. Palp short, with 5 segments, third palpomere without sensilla clavata subapically.

Thorax. Antepronotum well developed, with lobes meeting medially at anterior margin of scutum, with few ventrolateral antepronotals. Acrostichals small, scalpellate, in two rows starting some distance from antepronotum; dorsocentrals simple, uniserial; prealars few; supraalar absent. Scutellum apparently without setae.

Wing. Membrane without setae, with fine punctuation. Anal lobe reduced. With long costal extension. R2+3 running and ending close to R4+5; R4+5 ending distal to M3+4; FCu far distal to RM; Cu1 curved. Brachiolum with 1 seta, other veins bare. Squama bare. Sensilla campaniformia 3 above seta on brachiolum.

Legs. Tibial spurs normal, comb with few setae. Pseudospurs and sensilla chaetica absent, pulvilli vestigial.

Abdomen. Tergites with few setae, mostly in anterior band; sternites with few setae.

Hypopygium. Anal point long, triangular, with microtrichia, with or without strong setae at base, originating high on tergite IX or at posterior margin. Tergite IX with or without setae; laterosternite IX with few setae. Phallapodeme with aedeagal lobe well developed. Transverse sternapodeme arched or straight, without oral projections. Virga apparently consists of small spines. Gonocoxite long, without volsellae. Gonostylus without crista dorsalis, with long, weakly to strongly curved heel; megaseta normal.

Etymology

From Tupi Poxyaiba, ugly and Mberu, fly, meaning “the ugly fly”, referring to the spiny hypopygium of P. jamanximensis Andersen & Dantas, sp. nov. with a very large anal point. The name is masculine.

Poxyaibamberus jamanximensis Andersen & Dantas, sp. nov.

Figs 1A, B, 2, 3

Type locality

Brazil, Pará State, Itaituba, Jamanxim National Park; 05°41'58"S, 55°48'13"W; 170 m a.s.l.; 20 November 2017; Gilberto Nicácio leg.

Type specimen

Holotype male adult, slide-mounted in Euparal under six coverslips. Original label: “Brasil, PA, 20/11/2017, Floresta # 41, Shannon trap, Orthocladiinae, leg. G. Nicasio, ♂”. (INPA).

Diagnostic characters

The new species can easily be separated from P. ubajarensis Andersen & Dantas, sp. nov. by the shape of the gonostylus, as it has a rather narrow, weakly curved, tapering heel that is slightly longer than the gonostylus proper.

Description

Adult male (n = 1). Total length 2.17 mm. Wing length 1.23 mm. Total length / wing length 1.77. Wing length / length of profemur 2.51.

Coloration. Head, thorax, and legs light yellowish brown; abdomen pale yellowish brown. Wing hyaline.

Antenna (Fig. 1B). With 13 segments. AR = 0.94. Terminal flagellomere 340 µm long, with 21 µm long subapical seta.

Figure 1. 

Poxyaibamberus jamanximensis Andersen & Dantas, sp. nov. male (A, B) and P. ubajarensis Andersen & Dantas, sp. nov., male (C–F) A head, palpomere 4 and 5 not drawn B apex of ultimate flagellomere C palp D thorax E apex of fore tibia F apex of hind tibia.

Head (Fig. 1A). Inner verticals 5, outer verticals 4, postorbitals not discernable. Clypeus with 4 setae. Tentorium 99 µm long, 17 µm wide. Stipes 76 µm long. Anterior margin of cibarial pump slightly convex. Palp with palpomere 4 and 5 not measurable; palpomere 1–3 lengths (in µm) as: 14, 25, 38. Third palpomere without sensilla clavata.

Thorax. Antepronotum with 2 ventrolateral setae. Acrostichals about 8, scalpellate, in double row starting some distance from antepronotum; dorsocentrals 8, uniserial; prealars 3. Scutellum apparently without setae.

Wing (Fig. 2). VR = 1.33. Brachiolum with 1 seta, other veins and membrane bare. Squama bare. Costal extension 123 µm long.

Figure 2. 

Poxyaibamberus jamanximensis Andersen & Dantas, sp. nov. male. Wing.

Legs. Fore tibia with 33 µm long spur, mid tibia with 28 and 15 µm long spurs, hind tibia with 45 and 21 µm long spurs. Width at apex of fore tibia 27 µm, of mid tibia 28 µm, of hind tibia 29 µm. Hind tibia with comb of 4 bristles, longest apparently about 28 µm long. Lengths and proportions of legs as in Table 1.

Table 1.

Lengths (in µm) and proportions of legs of Poxyaibamberus jamanximensis Andersen & Dantas, sp. nov., male (n = 1).

Fe Ti ta1 ta2 ta3 ta4 ta5 LR BV SV BR
P1 490 596 547 270 155 82 49 0.918 2.941 1.985 2.10
P2 564 572 343 172 98 49 33 0.600 4.209 3.310 2.30
P3 605 629 417 212 163 82 41 0.662 3.311 2.961 2.40

Hypopygium (Fig. 3A–C). Anal point large, broadly triangular with rounded apex, starting high on tergite IX, with microtrichia and 12 strong setae in basal 1/3; 76 µm long, 39 µm wide near base, 18 µm wide medially. Laterosternite IX with 1 seta. Phallapodeme 76 µm long. Transverse sternapodeme arched, without oral projections, 62 µm long. Virga apparently consisting of field with small spines. Gonocoxite 203 µm long. Gonostylus straight, 104 µm long; heel weakly curved, tapering, 119 µm long; megaseta 7 µm long. HR = 1.95; HV = 2.09.

Figure 3. 

Poxyaibamberus jamanximensis Andersen & Dantas, sp. nov. male. A hypopygium, dorsal view B hypopygium with tergite IX and anal point removed, left dorsal aspect, right ventral aspect C gonostylus, ventral view.

Immatures and female. Larva, pupa, and female are unknown.

Etymology

The epithet, jamanximensis, is used as an adjective and meaning “from Jamanxim” in reference to the place of origin of the holotype.

Distribution

The species is only known from the type locality in Jamanxim National Park, Pará State, in northern Brazil, where it was collected in a Shannon trap placed near a lower-order stream. Only a single male was collected despite intensive sampling effort during the expedition with successive sampling using several types of traps. According to data from the National Institute for Space Research, the Jamanxim National Park ranks among the conservation units with the highest deforestation rates in the Amazon. This situation underscores the need to increase the knowledge of the biodiversity in this region to better understand the anthropogenic impacts on the biota and to utilize this knowledge as tools for developing conservation strategies. The discovery of a new genus of Chironomidae in the park highlights the importance of further research to understand and preserve its unique biodiversity.

Poxyaibamberus ubajarensis Andersen & Dantas, sp. nov.

Figs 1C–F, 4

Type locality

Brazil, Ceará State, Ubajara, Ubajara National Park, Cafundó waterfall, 03°50'13"S, 40°54'35"W, 805 m a.s.l., 25–26 February 2023, N. Hamada, J. Silva, J.M.C. Nascimento, G.P. Amorim Jr. leg.

Type specimen

Holotype male adult, slide-mounted in Euparal under five coverslips. Original label: “Brasil, CE, Ubajara, Parque Nacional de Ubajara, Cachoeira do Cafundó, 25-26/02/2023, light-trap, N. Hamada” (INPA).

Diagnostic characters

The new species can easily be separated from P. jamanximensis Andersen & Dantas, sp. nov. on the shape of the gonostylus, as it has a rather broad, curved, tapering heel that is slightly shorter than the gonostylus proper.

Description

Adult male (n = 1). Total length 2.02 mm. Wing length 1.13 mm. Total length / wing length 1.70. Wing length / length of profemur 2.38.

Coloration. Head, thorax, and legs light brown; abdomen pale brown. Wing hyaline.

Antenna. Antenna broken. With 44 µm long subapical setae.

Head. Inner verticals 5, outer verticals 3, postorbitals not discernable. Clypeus with 5 setae. Tentorium 90 µm long, 13 µm wide. Stipes not discernable. Anterior margin of cibarial pump slightly concave. Palp (Fig. 1C) with five segments; palpomere lengths (in µm): 17, 22, 35, 39, 37. Third palpomere without sensilla clavata.

Thorax (Fig. 1D). Antepronotum with 2 ventrolateral setae. Acrostichals very small, at least 4 in double row in anterior part of scutum; dorsocentrals 6, uniserial; prealars 3. Scutellum apparently without setae.

Wing. VR = 1.33. Brachiolum with 1 seta, other veins and membrane bare. Squama bare. Costal extension 113 µm long.

Legs. Fore tibia (Fig. 1E) with 30 µm long spur, mid tibia with 29 µm and 14 µm long spurs, hind tibia (Fig. 1F) with 35 µm and 18 µm long spurs. Width at apex of fore tibia 28 µm, of mid tibia 29 µm, of hind tibia 30 µm. Hind tibia with comb of 4 bristles, longest 22 µm long. Lengths and proportions of legs as in Table 2.

Table 2.

Lengths (in µm) and proportions of legs of Poxyaibamberus ubajarensis Andersen & Dantas, sp. nov., male (n = 1).

Fe Ti ta1 ta2 ta3 ta4 ta5 LR BV SV BR
P1 474 507
P2 507 515 310 155 98 53 45 0.602 3.795 3.297 1.92
P3 605 629

Hypopygium (Fig. 4A, B). Anal point large, narrowly triangular with rounded apex, projecting from posterior margin of tergite IX, with microtrichia, 55 µm long, 28 µm wide near base, 8 µm wide medially. Tergite IX with 9 setae medially. Laterosternite IX with 2 setae. Phallapodeme 68 µm long. Transverse sternapodeme straight, without oral projections, 41 µm long. Virga apparently consisting of balloon-shaped ball of lamella with small spines. Gonocoxite 168 µm long. Gonostylus 79 µm long; heel curved, tapering, 72 µm long, 24 µm wide medially; megaseta 8 µm long. HR = 2.13; HV = 2.56.

Figure 4. 

Poxyaibamberus ubajarensis Andersen & Dantas, sp. nov. male A hypopygium, dorsal view B hypopygium with tergite IX and anal point removed, left dorsal aspect, right ventral aspect.

Immatures and female. Larva, pupa, and female are unknown.

Etymology

The epithet, ubajarensis, is used as an adjective and meaning “from Ubajara” in reference to the place of origin of the holotype.

Distribution

The species is known only from the type locality in the Ubajara National Park in northeastern Brazil. The park covers an area of 6.288 ha and receives abundant rainfall, averaging 1.400 mm annually, while temperatures typically range between 22 °C and 26 °C. The vegetation in the park is characterized by its high diversity, with higher elevations adorned by lush humid forests (Figueiredo 1997; Souza 1997). Lower elevations feature semi-deciduous tropical rainforest on sloping areas and arboreal Caatinga (thorny deciduous forest) in the lower regions. The specimens were collected using a light trap placed about 20 m upstream of a waterfall (Fig. 5) at an elevation of 805 m a.s.l. At the sampling time, water temperature was 21.5 °C, pH was 7.05, and electric conductivity was 137.6 µS/cm.

Figure 5. 

The type locality of Poxyaibamberus ubajarensis Andersen & Dantas, sp. nov. in Ubajara National Park where the holotype was collected in a light trap near the Cafundó waterfall.

Discussion

In a consensus tree produced in the Bayesian analysis neighboring search, Poxyaibamberus falls out with Mesosmittia Brundin, 1956, Pseudosmittia Goetghebuer, 1932, Eretmoptera Kellog, 1900, Thalassosmittia Strenzke & Remmert, 1957, and Petalocladius Sublette & Wirth, 1972 (Fig. 6). Both Pseudosmittia, with more than 100 species, and Mesosmittia, with 18 species, are distributed in most parts of the world (Ferrington and Sæther 2011; Ashe and O’Connor 2012). The Neotropical species have been treated by Andersen et al. (2010) and Andersen and Mendes (2002). However, a position close to Thalassosmittia, Eretmoptera, and Petalocladius is notable. Thalassosmittia has 11 named species distributed in the Afrotropical, Nearctic, Neotropical, and Palaearctic regions (Ashe and O’Connor 2012). With few exceptions, Thalassosmittia representatives are marine shore dwellers (Andersen et al. 2013). However, the only Thalassosmittia described from the Neotropical Region is T. amazonica Andersen & Pinho, 2014, which was collected in the Amazon rainforest near Manaus (Andersen and Pinho 2014). Eretmoptera is a genus with two named species that are distributed in the Nearctic Region and in Antarctica. The adults are wingless. Eretmoptera browni Kellog, 1900 is distributed in California, while E. murphyi Schaeffer, 1914, was described from the island of South Georgia and is later introduced to Signy Island in the South Orkney Islands (Ashe and O’Connor 2012). The species is apparently parthenogenetic; the larva has a two-year life cycle and lives in damp moss and peat where they are thought to feed on decaying vegetation (Cranston 1985; Convey 1992). Petalocladius, with two included species, is distributed on the Caribbean Islands Jamaica and Hispaniola (Andersen et al. 2024). Both species have been collected in mountainous areas, and P. dominiensis Andersen & Baranov, 2024 was trapped close to a rather rapid, small river with rocky and stony substrates.

Figure 6. 

Position of Poxyaibamberus Andersen & Dantas, gen. nov. (marked in red) in the phylogenetic tree of Orthocladiinae, reconstructed with Bayesian inference (all posterior probabilities of the nodes are displayed, regardless of the value). This is a majority-rule consensus tree based on morphology only.

When posterior position probability mapping (“Rogue plots”) was applied, Poxyaibamberus was plotted in approximately 20% of the cases next to Stictocladius Edwards, 1931 and in approximately 20% of the cases next to Pseudosmittia / Eretmoptera (Fig. 7). While these were the most frequent positions within the generated trees, it plots near Eretmoptera in the consensus tree due to cumulatively more frequent occurrence (ca 40%) in or near this clade (Fig. 6). The position of Stictocladius in relation to Poxyaibamberus remains highly uncertain due to the lack of knowledge of the immature stages of the new genus. It is difficult to elaborate on the character distribution in the tree, as we have used an mcmc-based Bayesian approach, rather than maximum parsimony, which precludes more detailed analysis of the characters distribution. We have used the Bayesian approach, rather than maximum parsimony because of a deficiency in knowledge of many characters in the new genus. Thus, a more detailed analysis of the character distribution and sister relationships of Poxyaibamberus will have to wait until description of the additional life stages of the genus.

Figure 7. 

Rogue plot of the frequency of placement of Poxyaibamberus Andersen & Dantas, gen. nov. in the Bayesian tree from Fig. 6, based on the frequency of occurrence of Poxyaibamberus at a given node, based on the 50,001 most congruent trees from the Bayesian analysis of the genus’s morphology.

Acknowledgements

We are indebted to Jeane M.C. Nascimento, Jeferson O. da Silva, and Gilson P. Amorim Jr. for their invaluable support during the fieldwork in the Ubajara National Park, and to Gilberto Nicácio for providing us with the male from the Jamanxim National Park. NH received a CNPq research fellowship (process 308970/2019-5). GPSD is granted a postdoctoral fellowship by FAPEAM (process 01.02.016301.03271/2021-93). The following projects provided partial financial support: Sistemática integrada de insetos aquáticos na América do Sul (CNPq and Instituto Nacional de Pesquisas da Amazônia-INPA), and INCT ADAPTA II funded by CNPq (465540/2014–7) and FAPEAM (0621187/2017). Collection in Ubajara National Park was made under the SISBIO permit no. 82848-1. The University Museum of Bergen provided office space, microscope, and other equipment for TA.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

V. Baranov’s work is also funded by the State Agency of Innovation, within the Ramon y Cajal Program, grant number RYC2021-032144-I. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Vktor Baranov received funds supporting the payment of this publication fees through the CSIC Open Access Publication Support Initiative from the CSIC Unit of Information Resources for Research (URICI). CNPq research fellowship (Process 308970/2019-5) to N. Hamada. FAPEAM postdoctoral fellowship (Process 01.02.016301.03271/2021-93) to GPSD. CNPq and Instituto Nacional de Pesquisas da Amazônia-INPA funded the project “Sistemática integrada de insetos aquáticos na América do Sul”. CNPq (465540/2014–7) and FAPEAM (0621187/2017) funded the project “INCT ADAPTA II”. University Museum of Bergen, Bergen, Norway.

Author contributions

Trond Andersen – describing the new taxa, drawing the figures, preparation of manuscript, review of manuscript. Galileu P.S. Dantas – sorting and slide preparations, identification of specimens, preparation of manuscript. Viktor Baranov – phylogenetic analysis. Annui M. Sanz-laParra – photo, preparation of manuscript, review of manuscript. Humberto F. Mendes – preparation of manuscript. Neusa Hamada – organizing fieldwork, review of manuscript.

Author ORCIDs

Trond Andersen https://orcid.org/0000-0003-2201-1870

Galileu P. S. Dantas https://orcid.org/0000-0002-9155-533X

Viktor Baranov https://orcid.org/0000-0003-1893-3215

Annui M. Sanz-laParra https://orcid.org/0000-0003-0580-6890

Humberto F. Mendes https://orcid.org/0000-0002-8261-3665

Neusa Hamada https://orcid.org/0000-0002-3526-5426

Data availability

All of the data that support the findings of this study are available in the main text or in https://github.com/chironomus/Poxyaibamberus-.

References

  • Andersen T, Mendes HF (2002) Neotropical and Mexican Mesosmittia Brundin, with the description of four new species (Insecta, Diptera, Chironomidae). Spixiana 25: 141–155.
  • Andersen T, Mendes HF (2007) Five enigmatic new orthoclad genera from Brazil (Diptera: Chironomidae: Orthocladiinae). In: Andersen T (Ed.) Contributions to the systematics and ecology of aquatic Diptera. A tribute to Ole A. Sæther. The Caddis Press, Columbus, Ohio, 17–52.
  • Andersen T, Pinho LC (2014) A new Thalassosmittia Strenzke and Remmert, 1957 out of the sea: T. amazonica n. sp. from the Amazon rainforest, Brazil (Diptera: Chironomidae, Orthocladiinae). Chironomus 27(27): 25–30. https://doi.org/10.5324/cjcr.v0i27.1702
  • Andersen T, Sæther OA, Cranston PS, Epler JH (2013) 9. The larvae of Orthocladiinae (Diptera: Chironomidae) of the Holarctic Region — keys and diagnoses. In: Andersen T, Cranston PS, Epler JH (Eds) Chironomidae (Diptera) of the Holarctic Region. Keys and diagnoses — larvae. Insect Systematics & Evolution, (Supplement 66), 189–386.
  • Andersen T, Mendes HF, Pinho LC (2015b) Mariambera, a new genus of Orthocladiinae from Brazil (Insecta: Diptera, Chironomidae). Studies on Neotropical Fauna and Environment 50(1): 24–30. https://doi.org/10.1080/01650521.2014.991589
  • Andersen T, Höcherl A, Sanz-laParra AM, Bastardo RH, Baranov V (2024) A new species of Petalocladius Sublette & Wirth, 1972 (Diptera, Chironomidae, Orthocladiinae) from the Dominican Republic, with an emended generic diagnosis. Chironomus. Journal of Chironomidae Research 38: 4–13.
  • Ashe P, O’Connor JP (2012) A world catalogue of Chironomidae (Diptera). Part 2. Orthocladiinae (Section A & B). Irish Biogeographical Society and National Museum of Ireland, Dublin, 968 pp.
  • Convey P (1992) Aspects of the biology of the midge, Eretmoptera murphyi Schaeffer (Diptera: Chironomidae), introduced to Signy Island, maritime Antarctic. Polar Biology 12(6–7): 653–657. https://doi.org/10.1007/BF00236988
  • Cranston PS (1985) Eretmoptera murphyi Schaeffer (Diptera: Chironomidae), an apparently parthenogenetic Antarctic midge. British Antarctic Survey Bulletin 66: 35–45.
  • Ferrington Jr LC, Sæther OA (2011) A revision of the genera Pseudosmittia Edwards, 1932, Allocladius Kieffer, 1913, and Hydrosmittia gen. n. (Diptera: Chironomidae, Orthocladiinae). Zootaxa 2849(1): 1–314. https://doi.org/10.11646/zootaxa.2849.1.1
  • Figueiredo MA (1997) Unidades fitoecológicas. In: Iplance (Ed.) Atlas do Estado do Cerá. Fundaçào Instituto de Planejamento do Ceará, Governo do Estado do Ceará, Seplan, Fortaleza, 28–29.
  • Fittkau JE (1974) Ichthyocladius n. gen., eine neotropische Gattung der Orthocladiinae (Chironomidae, Diptera) deren Larven epizoisch auf Welsen (Astroblepidae und Loricariidae) leben. Entomologisk Tidskrift (Supplement 95): 91–106.
  • Klopfstein S, Spasojevic T (2019) Illustrating phylogenetic placement of fossils using RoguePlots: An example from ichneumonid parasitoid wasps (Hymenoptera, Ichneumonidae) and an extensive morphological matrix. PLoS ONE 14(4): e0212942. https://doi.org/10.1371/journal.pone.0212942
  • Mendes HF, Andersen T (2008) A review of Antillocladius Sæther and Litocladius Mendes, Andersen et Sæther, with the description of two new Neotropical genera (Diptera, Chironomidae, Orthocladiinae). Zootaxa 1887(1): 1–75. https://doi.org/10.11646/zootaxa.1887.1.1
  • Mendes HF, Andersen T, Sæther OA (2004) New species of Ichtyocladius Fittkau, a member of the Corynoneura-group (Diptera: Chironomidae: Orthocladiinae) with a review of the genus. Studies on Neotropical Fauna and Environment 39(1): 15–35. https://doi.org/10.1080/01650520412331270936
  • Mendes HF, Sæther OA, Andrade-Morraye M (2005) Gynocladius scalpellosus n. gen., n. sp. from Brazil (Diptera: Chironomidae: Orthocladiinae). Zootaxa 979(1): 1–12. https://doi.org/10.11646/zootaxa.979.1.1
  • Mendes HF, Andersen T, Pinho LC (2013) Uirassubrillia beckeri gen. n., sp. n. (Diptera: Chironomidae, Orthocladiinae) from northeastern Brazil. Zootaxa 3716(1): 65–74. https://doi.org/10.11646/zootaxa.3716.1.5
  • Page RDM (2001) NDE: NEXUS data editor 0.5. 0. University of Glasgow, Glasgow.
  • Ronquist F, Teslenko M, van der Mark P, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61(3): 539–542. https://doi.org/10.1093/sysbio/sys029
  • Sæther OA (1969) Some Nearctic Podonominae, Diamesinae and Orthocladiinae (Diptera: Chironomidae). Bulletin - Fisheries Research Board of Canada 107: 1–154.
  • Sæther OA (1980) Glossary of Chironomid morphology terminology (Diptera: Chironomidae). Entomologica Scandinavica (Supplement 14): 1–51.
  • Schaeffer C (1914) Collembola, Siphonaptera, Diptera, and Coleoptera of the South Georgia Expedition. Science Bulletin. Museum of the Brooklyn Institute of Arts and Sciences 2(4): 90–94.
  • Souza MJN (1997) Geomorfologia. In: Iplance (Ed.) Atlas do Estado do Cerá. Findaçâo Instituto de Planejamento do Cerá, Governo do Estado do Cerá, Seplan, Fortaleza, 18–19.
  • Wiedenbrug S, Trivinho-Strixino S (2009) Ubatubaneura, a new genus of the Corynoneura group (Diptera: Chironomidae: Orthocladiinae) from the Brazilian Atlantic Forest. Zootaxa 1993(1): 41–52. https://doi.org/10.11646/zootaxa.1993.1.4
login to comment