John H. Epler 461 Tiger Hammock Road, Crawfordville, Florida, 32327

CHIRONOMUS Journal of Chironomidae Research No. 30, 2017: 4-18. Current Research.

An annotated preliminary list of the Chironomidae (Diptera) of Zurquí, Costa Rica

John H. Epler

461 Tiger Hammock Road, Crawfordville, Florida, 32327, U.S.A. Email: [email protected]

Abstract to October 2013. The 150 by 266 m site, at an el- evation of ~1600 m, is mostly cloud forest, with An annotated list of the species of Chironomidae adjacent small pastures; the site has one permanent found at a four-hectare site, mostly cloud forest, in and one temporary stream, located in heavily for- Costa Rica is presented. A total of 137 species, 98 ested ravines. of them undescribed, in 63 genera (17 apparently new), were found. Collecting methods included two malaise traps run continuously and additional traps run three days Introduction each month: three additional malaise traps, several The tropics have long been known as areas of great emergence traps (over leaf litter; over dry branch- biodiversity (e.g. Erwin 1982), but our knowledge es; over vegetation; over stagnant water; over run- of many insect groups there remains poor. The two ning water), CDC light traps, bucket light traps, volume “Manual of Central American Diptera” yellow pan traps, flight intercept traps and mercury (Brown et al. 2009, 2010) provided the first modern vapor light traps. Some specimens were collected tools to analyze the diversity of one of the largest by sweeping and by hand. orders of insects, the Diptera (two-winged flies) of Samples were sorted and prepared by technicians the northern portion of the Neotropics; Spies et al. at the Instituto Nacional de Biodiversidad (INBio). (2009) covered the Chironomidae. These volumes Given the difficulties of female identification, for offered the first family level identification keys for the most part only males were removed from the Central America (the first for any portion of the samples for identification. Thus parthenogenetic Neotropics) and also allow identification to the ge- species, such as Phytotelmatocladius delarosai neric level of the described Central American Dip- Epler, known from the USA (Florida) to Argentina tera. However, these volumes are just scratching (Epler 2010, Siri & Donato 2014) may have been the surface of Neotropical fly diversity. At both the missed. genus and especially the species level, a tremen- dous amount of work remains to be done. Results Art Borkent and Brian Brown conceived and de- A total of 2,120 specimens were examined. Four veloped a survey project for identifying all the subfamilies, 63 genera and 137 species were repre- Diptera species of a small area in Costa Rica. The sented (Appendix 1). The majority of species (98), study, patterned after an “All Taxa Biotic Invento- and 17 new genera, are apparently undescribed and ry” (ATBI), but devoted entirely to the order Dip- were given letter/number designators. It is antici- tera, was called the Zurquí All Diptera Biodiversity pated that a great deal of time will be necessary to Inventory (ZADBI). Fifty-nine collaborators took describe all these new taxa. Workers whose work part in the project; I was fortunate to work with the may involve some of these taxa are welcome to Chironomidae. Borkent & Brown (2015) provided contact me with ideas for producing descriptions. an overview of the project and its protocol. Funding sources are also being sought. As will be seen below, a large number of unde- All but one of the putatively new genera were or- scribed genera and species were found, as well as thoclads and are listed as “CHIRORTH-n”, refer- many new records for Costa Rica. This paper is ring to “Chironomidae: Orthocladiinae” followed also meant to update the generic synopses provid- by a number; e.g., “CHIRORTH-1 sp. ZUR-1”. ed by Spies et al. (2009). The single CHIRCHIR-1 is a damaged specimen of a Chironominae, and may represent a known Material and methods genus, but its combination of available characters The majority of material came from the main site does not allow generic placement. Undescribed at Zurquí de Moravia (hereafter Zurquí), located species are listed with “ZUR-n”, referring to new at 10.047313°N, 84.008457°W, in San José Prov- taxa from the Zurquí project (I also have numer- ince, Costa Rica, collected from September 2012 ous other undescribed species from Costa Rica,

4 collected over the past several decades, that have a single specimen. A key character for this species been given different letter/number designators; was the absence of acrostichal setae. The Zurquí with the possible exception of a single species material (41 specimens) all had some acrostichal of Parametriocnemus, none of these other unde- setae; otherwise the specimens appear to be B. scribed taxa were found in the Zurquí material). A psilacrus. The type was not available for examina- list of the taxa collected is presented in Appendix tion. 1. An annotated accounting of these taxa follows Although a widespread and speciose genus, few below; any mention of morphological characters Bryophaenocladius species are known from the is not to be taken as a description of a taxon. Note Neotropics. Wang et al. (2006) and Donato (2011) also that the relative numbers or percentages given provided the most recent treatments of the genus for species collected are not representative of the for the area. It is possible that some of the taxa that total catch, and pertain just to the material that was I’ve grouped here may belong elsewhere. selected to be mounted. Compterosmittia - Three species were collected; PODONOMINAE the most common (104 specimens) was the unde- Parochlus - A single taxon belonging to the Pa- scribed C. sp. ZUR-1, followed by C. nerius (43 rochlus araucanus group (Brundin 1966) was col- specimens). A single specimen of C. sp. ZUR-2 lected. Because pupae are necessary for species was collected. The genus was most recently re- delimitation in this group, species-level identifica- viewed, for taxa from the area, by Mendes et al. tion of this taxon will have to wait until pupae are (2004). collected and associated. Corynoneura - Two undescribed species were col- TANYPODINAE lected in addition to two described species: C. ferelobata was described from Guatemala by Sublette Procladius - To date, only one species of the sub- & Sasa (1994); C. guanacaste was described from genus P. (Procladius) has been reported from the northern Costa Rica by Wiedenbrug et al. (2012). Neotropics: P. mozambique Roback, from Colom- bia (Roback 1982b). The single species of Procla- Cricotopus - About 15 species are recorded or de- dius collected at Zurquí, P. (Procladius) sp. ZUR- scribed from the Central American-Caribbean area 1, more closely resembles P. (P.) paludicola Skuse, (Spies & Reiss 1996). All eight of the Cricotopus an Australian species (Roback 1982a), but differs species that were collected at Zurquí appear to be in coloration and morphology. undescribed; all belong with the subgenus C. (Cri- cotopus). Thienemannimyia - Thienemannimyia (Thiene- mannimyia) sp. ZUR-1 may be the species referred Diplosmittia - Two undescribed species were col- to by Watson & Heyn (1993) as Thienemannimyia lected, in addition to D. forficata, described from nr barberi (Coq). It resembles T. barberi (Coquil- La Selva, Costa Rica, by Andersen (1996). The lett), a western US species, but the wing pattern genus has recently been reviewed by Pinho et al. appears slightly different; comparison of more (2009) and Wiedenbrug & Silva (2016) added an Costa Rican and Nearctic material is necessary. additional species from the Dominican Republic. The single specimen of D. sp. ZUR-1lacks anten- Zavrelimyia - Zavrelimyia (Paramerina) fasciata, nae and bears a well developed inferior volsella; it as Paramerina fasciata Sublette & Sasa, was de- may belong elsewhere but is placed in Diplosmit- scribed from Guatemala (Sublette & Sasa 1994); tia until more complete material can be collected. it is widespread in Costa Rica and was the most Diplosmittia sp. ZUR-2 resembles D. beluina common tanypod collected in the study. Andersen (described from La Selva, Costa Rica) ORTHOCLADIINAE but has 13 antennal flagellomeres, with an AR of 3.0-3.4, and a more sharply pointed anal point; D. Antillocladius - Three species were collected; A. beluina has 10 flagellomeres and an AR of 0.62. pluspilalus was the most common (59 specimens). Both taxa have a long costal extension. The two The genus has most recently been reviewed by taxa may be conspecific but more material is nec- Mendes & Andersen (2008) and Mendes et al. essary to determine antennal vaiability. (2011). Eukiefferiella - Three undescribed species that fit Bryophaenocladius - Of the five Bryophaeno- Eukiefferiella were collected. No described spe- cladius species collected, four are undescribed. cies are known from the Neotropics; Watson & The fifth species is listed here as B. cf. psilacrus. Heyn (1993) recorded larvae from Costa Rica. Bryophaenocladius psilacrus was described from South Carolina, USA, by Sæther (1982), based on

5 Gravatamberus - The genus was established by May, with fewer setae present in those wing areas Mendes & Andersen (2008); they included five from May (one specimen) through October. species found in Brazil, Chile, Costa Rica, Gua- Irisobrillia - Eight specimens of the sole species of temala, Mexico and Venezuela. Gravatamberus the genus, I. longistyla, were collected. curtus Mendes & Andersen was described from material (three specimens) from Mexico and Krenosmittia - Two specimens were collected of a northwestern Costa Rica; G. guatemaltecus was species that appears to belong with Krenosmittia. described from a single specimen from Guatemala. This would be the first record for the genus from The two species were diagnosed by the number of the Neotropics. Examination of more material is setae in cell m proximal to RM, number of setae on necessary. the subcosta, the length of the costal extension and Limnophyes - Limnophyes was one of the more the AR: G. curtus has <10 setae in cell m proxi- common of the small orthoclad taxa collected mal to RM (G. guatemaltecus >10); subcosta with (~8% of total mounted catch). Two species were 6–15 setae (G. guatemaltecus 16), costal extension collected: L. mariae (38 specimens) and the much 68-86 µm (G. guatemaltecus 161); AR 0.64–0.69 more common L. guatemalensis (142 specimens), (G. guatemaltecus 0.26). both described from Guatemala by Sublette & Eighteen males of Gravatamberus were collected; Sasa (1994). unfortunately, antennae were missing on all of Lipurometriocnemus - Four species were col- them. All had a costal extension, ranging from 128 lected, three of them undescribed. Lipurometrioc- to 200 µm (Table 1), greater than that recorded for nemus was first described from the British West In- G. curtus. However, the range of setae in cell m dies by Sæther (1981), with L. glabalus as the sole proximal to RM ranged from 0 to 24 in the Zur- species. Sæther (1982) then described L. vixloba- qui material, regardless of costal extension length, tus from South Carolina, USA. The two species suggesting that this character does not aid in dis- were supposedly separated by the absence of setae tinguishing G. curtus from G. guatemaltecus. All on R , a weak inferior volsella and a “slight hump Gravatamberus collected in this study are consid- 1 on tergite IX representing a reduced anal point” in ered to represent G. guatemaltecus. L. vixlobatus. Cranston & Oliver (1988) reported Data also indicate that the midges had more se- L. vixlobatus from the Yukon Territory of Canada, tae in cell m and on the subcosta from February to and examined paratypes of L. glabalus and a sin-

Table 1. Selected measurements of Gravatamberus guatemalticus Specimen # Month collected cell m setae subcosta setae costal extension, µm 4399193 OCT 0 5 175 4401710 JUN-JUL 1 4 195 4401650 JUN-JUL 0 6 200 4401625 OCT 0 5 155 4400249 JUN 1 5 180 4405861 MAY 1 5 153 4401624 OCT 0 9 200 4400246 JUN 1 7 188 4399225 JUN 0 5 155 4399224 JUN 9 5 175 4406521 APR 18 12 163 4370228 MAY 17 17 153 4363952 APR 24 14 128 4363944 FEB 11 7 188 4363957 APR 19 11 133 4399449 MAY 15 10 165 4406482 APR 13 6 150 4406453 MAY 16 10 138

6 gle specimen (which they thought to also represent from a single male specimen from Panama (Sæther L. glabalus) from Braulio Carillo National Park in 1985), was separable from M. patrihortae only Costa Rica (the Zurquí site is adjacent to a portion by the length of the costal extension (116 µm in of the huge park). They noted that the two species M. truncata; 8–62 µm in M. patrihortae). Costal were difficult to separate because of variation in extensions in Zurquí material ran from 50 to 140 the supposedly diagnostic characters, but gave no µm, indicating a wide range of lengths that would reason why they assigned the Yukon specimen to include the single measurement from the Panama vixlobatus, which following its description should specimen. I consider M. truncata to be a junior lack any setae on R1, instead of assigning it to L. synonym of M. patrihortae. glabalus. I noted such variation in the volsellae and Metriocnemus - Three species, two undescribed, “anal hump” in the Zurquí material I’ve designated were collected. The named species, Metriocnemus as L. glabalus; there was variation in the length/ costatus, was described from Guatemala by Sub- breadth of the gonostylus but much appeared due lette & Sasa (1994). Their description stated “geni- to the viewing angle. I saw no specimens in which talia with a midventral notched, somewhat quad- R was devoid of setae and considered all material 1 rangular plate”. One of the Metriocnemus species I put in this group to be L. glabalus (63 specimens). collected, initially designated M. sp. ZUR-2, was My L. sp. ZUR–1(eight specimens) has genitalia very similar to the description and illustrations for similar to those of L. glabalus but has a very low M. costatus (Sublette & Sasa 1994: fig. 88) except AR (~0.32–0.58) compared to what is being called it apparently lacked this plate. I was able to exam- L. glabalus today (following Cranston & Oliver ine 6 paratypes of M. costatus from the Sublette 1988 and Andersen et al. 2016). In Sæther’s (1981) Collection, located at the University of Minnesota, original description of L. glabalus, all of the type St. Paul, MN. The specimens were all excessively material lacked antennae, but it is now apparently squashed (typical of Sasa mounts) and the quad- assumed that the species has an AR similar to that rangular plate was apparent. I then remounted the of L. vixlobatus – around 1.60–1.75. Zurquí mate- abdomen/genitalia of one of the Zurquí M. sp. rial of L. glabalus had ARs > 1.70. ZUR-2 specimens and excessively squashed it – the quadrangular plate appeared! This plate is nor- My L. sp. ZUR–2 (36 specimens) has an AR around mally oriented in a dorsal-ventral manner and is 1.76–1.85 and genitalia very similar to those of L. visible as a thin sclerotized line when viewed in glabalus, except the proximal portion of the gono- a typical genitalia mount that has not been exces- stylus bears a hump that separates it from that of sively flattened. The structure may represent the L. glabalus. Further work (i.e., measurements of aedeagus or an ejaculator (see Tuxen 1970: fig. other body parts, etc.) may show this taxon may 160). also belong with L. glabalus. Nanocladius - Eight specimens of a single un- My L. sp. ZUR–3 (two specimens) has an AR of described species of Nanocladius (Nanocladius) about 1.0 or less and has a distinctively pronounced were collected. The genus is poorly known from inferior volsella. Andersen et al. (2016) described the Neotropics, with only three species described two new Lipurometriocnemus species from Brazil from the region to date (Epler 1986; Wiedenbrug and provided a key for the known species. Their L. & Silva (2013). biancae has a pronounced inferior volsella, but not the same as L. sp. ZUR–3, and an AR of 1.29-1.45. Onconeura - One undescribed species (six speci- Their L. amazonicus (AR 1.23–1.45) is very simi- mens) of Onconeura was collected. This would be lar to L. glabalus, but has fewer dorsocentral setae. the second species known from Costa Rica; Spies & Reiss 1996) recorded O. semifimbriata (Sæther) Litocladius - A single species (34 specimens), L. from the country. The most recent papers on On- chavarriai, was collected. coneura are Donato et al. (2012) and Wiedenbrug Lopescladius - Two undescribed species, both et al. (2009). belonging with the subgenus L. (Cordiella), were Parakiefferiella - The genus is unrecorded from collected (seven of L. sp. ZUR-1; two of L. sp. Central America; Wiedenbrug & Andersen (2002) ZUR-2). described several new species from South America Mesosmittia - The widespread M. patrihortae and provided a review of the genus. Two speci- (22 specimens) and one undescribed species (one mens of an undescribed species were collected. specimen) were collected. The genus was most re- Parametriocnemus - Four species were collected; cently reviewed by Andersen & Mendes (2002). all appear to be undescribed. Parametriocnemus They noted that M. truncata Sæther, described sp. ZUR-1 and P. sp. ZUR-2 resemble the Nearc-

7 tic P. lundbeckii (Johannsen) but have a lower AR; the most common (75 mounted specimens), while P. sp. ZUR-1 was 0.67–0.72; P. sp. ZUR-2 was 39 specimens of the widespread P. forcipata were 0.52-0.68 (for P. lundbeckii see below). Both may collected. It should be noted that, pending further represent a single species; more measurements of investigation, some of the putative new orthoclad other body parts are necessary. genera found during this study may belong with Pseudosmittia. Ferrington & Sæther (2011) pro- Sublette (1967: 539) gave an AR of 1.40 based on vided the most recent review of the genus. a specimen (a “metatype”) from the type series; he also gave ARs of four Johannsen-determined Stictocladius - One specimen of an undescribed specimens that ranged from 1.26–1.60, mean 1.36. species was collected. In the Western Hemisphere, Sæther (1969) gave a range of 0.90-1.30, mean this genus has been mostly recorded from Argen- 1.12, for the AR of P. lundbeckii. tina, Chile and Peru, but has recently been found in North America (Sæther & Cranston 2012). This Sublette & Sasa (1994) recorded larval, pupal and would be the first record of the genus from Central adult P. lundbeckii from Guatemala. I examined 4 America. The new Costa Rican species has clear females and one female pupal exuviae from this wings and a very low AR of 0.21. Guatemalan material (no males were available). None of these specimens could be reliably identi- Synorthocladius - Two specimens of an unde- fied as P. lundbeckii. scribed species were collected. Watson & Heyn (1993) recorded “Synorthocladius sp.” from Costa Epler (2001: 7.117) discussed a Parametriocnemus Rica. from Great Smoky National Park that had genitalia similar to P. lundbeckii but had an AR of 0.40. Two The following orthoclad taxa appear to represent other specimens resembling P. lundbeckii from the undescribed genera. These taxa are discussed be- same area have an AR of around 0.84, and I’ve low with mention of morphological characters also examined P. lundbeckii from the same area in order to inform other workers that may have with “typical” ARs (~1.40). It appears obvious that similar taxa and may wish to collaborate or bor- a thorough study of P. lundbeckii is necessary to row material. Please note again that any mention determine just what that species is. of morphological characters is not to be taken as a description of a taxon. According to my notes, P. sp. ZUR-4 resembles a species I have seen from northwestern Costa Rica CHIRORTH-1 - A small species, represented by that I had designated P. sp. CR-3. Those speci- one specimen with bare eyes; 11 antennomeres; mens, and other Costa Rican Parametriocnemus minute scalpellate (?) acrostichals; wing with species, are out on loan and were not available for strong punctation and bare squama; small rounded comparison. All of the Zurquí Parametriocnemus anal point; and each tarsal claw with 3 teeth. specimens had an AR below 1.00. CHIRORTH-2 - A small species with bare eyes; Paraphaenocladius - Paraphaenocladius exagi- 13 antennomeres with low (0.43) AR; elongate tans longipes was one of the most common or- thorax with a few decumbent acrostichals; 0-1 thoclads collected, with 113 specimens examined. squamal setae, with long costal extension; abdo- The subspecies was described from Costa Rica men with weak bands; no virga, no anal point. (type locality, Cacao (a volcano) in Guanacaste CHIRORTH-3 - A small species with pubescent Province), St. Vincent and Trinidad (Sæther & eyes; low AR (0.50); well developed scale-like vir- Wang 1995). ga; and a short, rounded anal point. Fifteen speci- Pseudorthocladius - Two specimens of an appar- mens were collected. ently undescribed species were collected. Until CHIRORTH-4 - There appear to be two species this collection, Pseudorthocladius was not re- in this putative new genus, which differ from each corded from the Neotropical Region. The Zurquí other by having a radically different virga. The species appears similar to P. clavatosus Sæther & head has a U-shaped frons as in Jururumberus Sublette, but has a smaller inferior volsella. Pseu- Mendes & Andersen (Mendes & Andersen 2013). dorthocladius clavatosus was described from two This small-bodied genus also has bare eyes; 8/9 males and a female from South Carolina, U.S.A. antennomeres; no squamal setae; R ends just be- (Sæther & Sublette 1983); comparison with type fore fCu and no anal point. Numerous specimens material will be necessary to determine if the Zur- were collected. quí specimens represent a new species. CHIRORTH-5 - A genus similar to Pseudosmit- Pseudosmittia - Three species were collected, one tia, represented by four specimens. It features undescribed. Pseudosmittia windwardensis was

8 antennae with 13 flagellomeres; low AR (0.55); palpomere; apparently no acrostichals; bare squa- weak decumbent acrostichals; one squamal seta; ma; elongate virga; and broadly spatulate anal wing veins without setae except near base of R; point. Three specimens were collected. long costal extension; no pulvilli and deeply bifid CHIRORTH-14 - One specimen of this taxon was tarsal claws. collected. It has bare eyes; 13 flagellomeres, with CHIRORTH-6 - This taxon, represented by four low AR (0.28) and a large seta on the last antenno- specimens, resembles Saetheriella Halvorsen, mere; no acrostichals; no squamals; a moderately in having reniform, protruding hairy eyes and a long bare anal point and a well developed inferior rounded to triangular anal point. However, it has volsella. an extended costa; well developed pulvilli; a well CHIRORTH-15 - A series of 10 specimens rep- developed, clear, scutal tubercle; lacks a virga; and resents this genus, which has bare eyes; short pal- has superior and inferior volsellae. pomeres; two acrostichals at mid-scutum; long CHIRORTH-7 - This genus has pubescent eyes; pulvilli; and no anal point. All specimens have lost no observable acrostichal or squamal setae; a very their antennae. long costal extension; a scale-like virga; and a CHIRORTH-16 - A single specimen of this midge short rounded anal point. Three specimens were was collected. It somewhat resembles Pseudosmit- collected. tia, but has hairy eyes; no acrostichal or squamal CHIRORTH-8 - A single specimen of this taxon setae; a long costal extension; weak virga; small was collected. It has eyes with very short pubes- inferior volsellae; arcuate gonostyli; and a short cence; antennae with 13 flagellomeres and very triangular/conical anal point lacking any large se- low AR (0.19); no apparent acrostichals; no squa- tae. mals, no pulvilli; apically bifid claws; no volsella CHIRONOMINAE and a gonostylus that is widened preapically. Beardius - Two species were collected; B. triangu- CHIRORTH-9 - Represented by two specimens, latus was described from Costa Rica by Andersen this genus has pubescent eyes, with pseudo-ocelli & Sæther (1996). One specimen of an undescribed present; a grossly enlarged globose palpomere 3; species was collected. The genus was recently re- numerous scalpellate acrostichals; a small patch vised by Pinho et al. (2013). of short setae lateral to and above the apex of the antepronotal lobes; no squamals; a long costal ex- Caladomyia - Two undescribed species were col- tension; very recurved Cu1 and a Mesosmittia-like lected; one specimen of one species, two of the anal point. other. The genus was recently reviewed by Trivin- ho-Strixino (2012). CHIRORTH-10 - A single specimen of this moderately large midge was collected. It has 13 flagel- Cryptochironomus - Four specimens of an unde- lomeres; bare eyes; reduced 4th and 5th palpomeres; scribed Cryptochironomus were collected. about 20 scalpellate acrostichals that originate at Dicrotendipes - Fourteen specimens of an unde- about 1/3 the length of the scutum; finely punc- scribed Dicrotendipes were collected. The species tate wing membrane; no squamals; sinuate Cu1; possesses an apically bifid inferior volsella. Al- no anal point but a Mesosmittia-like median hump; though such a volsella is known from Afrotropical abdominal tergites with dark posterior bands; and and Palaearctic species, as well as several Amazo- a well developed inferior volsella with a median nian species (Epler 1988), this is the farthest north swelling posterior and ventral to it. I have seen such a species in the Western Hemi- CHIRORTH-11 - Two specimens were collected; sphere. both have lost their antennae. Eyes are bare; the Einfeldia - Five specimens of an undescribed Ein- acrostichals are absent or very minute; a weak feldia were collected. The taxonomy of Einfeldia virga is present; there are no volsellae present; and has been confused, with some species being allo- the gonostylus lacks a megaseta. cated to other genera (Epler et al. 2013; Cranston CHIRORTH-12 - A single specimen that lost its et al. 2016). I examined the holotype of E. atitlan- antennae was collected. It has hairy eyes; four pal- ensis Sublette & Sasa, described from Guatemala pomeres; no squamal setae; apparently no acros- (Sublette & Sasa 1994); this new species differs tichals; a virga; Mesosmittia-like anal point; and a having a larger digitus of the superior volsella, a small inferior volsella. thinner gonostylus and darkened apices of the leg segments. CHIRORTH-13 - This taxon has bare eyes; one

9 Endotribelos - Two described species and one un- and confirmed the genus’s position within the tribe described species were collected. Fourteen speci- Tanytarsini. mens of E. albatum and 40 (including two inter- Nilothauma - Two undescribed species were col- sex specimens) of E. grodhausi were examined; lected. The genus was most recently reviewed by seven specimens of the undescribed species were Mendes & Andersen (2009). collected. The undescribed species has a long thin anal point and appears similar to, but lighter than, Parachironomus - Four specimens of an unde- an undescribed species from La Selva, of which I scribed Parachironomus were collected. The Ne- have several reared associations. otropical Parachironomus were documented by Spies et al. (1994). Endotribelos was established by Grodhaus (1987) for Tendipes (Tribelos) hesperia Sublette, 1960 Phaenopsectra - A single specimen of an unde- (which thus became Endotribelos hesperium (Sub- scribed Phaenopsectra was collected. lette)), a southern US species whose larva has an Polypedilum - A very speciose genus world-wide, odd number of teeth on the mentum and a large Polypedilum was the most abundant chironomine gap (diastema) between the molar area and the genus collected in the study, with one undescribed proximal inner tooth of the mandible. Subsequent- species of P. (Pentapedilum), one described and ly, Sublette & Sasa (1994) described two species three undescribed P. (Polypedilum), three de- from Guatemala, E. albatum and E. grodhausi; the scribed and one undescribed P. (Tripodura) and larva of E. grodhausi has an even number of men- one undescribed species of P. (Uresipedilum). tal teeth and lacks the mandibular diastema (the larva of E. albatum remains unknown). Since then, The undescribed P. (Pentapedilum) species was several more species have been described from the most abundant Polypedilum collected, with 88 South America (Roque & Trivinho-Strixino 2008 specimens mounted. Mendes, Andersen & Jocqué and Trivinho-Strixino & Pepenelli 2015), the lar- (2011) described P. (Polypedilum) panacu from vae of which display a variety of those character bromeliad phytotelmata in Honduras. Of the three states. See also Epler et al. (2013). described P. (Tripodura) collected, two were described by Sublette & Sasa (1994) from Guatema- Trivinho-Strixino & Pepenelli (2015) provided la. One of these, P. (Tripodura) luteopedis was the keys for the males and known larvae of Endotribe- second most abundant Polypedilum collected, with los. However, male E. grodhausi will not key 75 specimens mounted; the widespread P. (T.) api- correctly because couplet 4 gives one the choice catum was described from New Mexico, U.S.A., of “base of superior volsella setose” or “base of by Townes (1945) and was also recorded from superior volsella bare”. Choosing “bare” eventu- Guatemala by Sublette & Sasa (1994). ally leads to E. grodhausi at couplet 14. However, the base of the superior volsella of E. grodhausi Rheotanytarsus - One undescribed species, two is clothed, ventrally and dorsally, with fine se- described species and two tentatively identified tae. Thus, E. grodhausi will key to E. jaragua described species were collected. Rheotanytarsus Trivinho-Strixino & Pepenelli in couplet 5, from guancastensis, R. scutulatus and R. subtilis were which it may be separated by the darker thorax of described from northwestern Costa Rica by Ky- E. grodhausi (preepisternum and adjacent sclerites erematen & Andersen (2002); R. contrerasi was are dark; light in E. jaragua). Sasa and Sublette’s described from Mexico by Andersen & Sæther in illustration of the volsella (Sublette & Sasa 1994: Kyerematen et al. (2000). Our specimens of R. cf. fig. 141) does not indicate the presence of setae, contrerasi and R. cf. guanacastensis differ slightly which apparently led Trivinho-Strixino & Pep- from their original descriptions and must be com- enelli (2015) to assume the structure was bare. pared with type material for more positive identi- Sublette & Sasa (1994: fig. 135) likewise illustrat- fication. ed the base of the superior volsella of E. albatum Riethia - Fifteen male specimens of a genus from as bare, when it also is clothed with fine setae. I the putative tribe Pseudochironomini were collect- examined two paratypes of E. albatum and eight ed; this genus is almost certainly Riethia, but there paratypes of E. grodhausi. is a small chance they could be Manoa. Males of Nandeva - Two specimens of N. latiloba were the two genera cannot at this time be separated collected; the species was described from Brazil utilizing morphological characters; only females by Sæther and Roque (2004). This is the first re- and pupae may be separated at the generic level cord of the genus from Costa Rica. Andersen et al. (Jacobsen & Perry 2002; Trivinho-Strixino et al. (2011) also recorded the species from Venezuela 2009). I’m calling the Zurquí specimens Riethia based on their close resemblance to R. truncato-

10 caudata (Edwards) as redescribed in Trivinho- (2007) from Brazil, but appears a bit different; T. Strixino et al. (2009) and R. manauara Neubern, sp. ZUR-8 (one specimen) is also similar to T. cf. Trivinho-Strixino & Silva (Neubern et al. 2011); caipira but is much smaller. Tanytarsus sp. ZUR- the medial apices of the inferior volsellae of all 5 (16 specimens) is similar to T. cotopaxi Giłka three species have large, flattened, apically pecti- & Zakrzewska, described from Ecuador (Giłka nate setae directed mediad (and dorsad in R. trun- & Zakrzewska 2013). The majority of specimens catocaudata). Such setae are unknown in Manoa. were T. sp. ZUR-1 (47 specimens), including an Note that the legs of the Zurquí specimens are un- intersex specimen; 23 specimens of T. sp. ZUR-2; banded and there are other differences, especially 34 specimens of T. sp. ZUR-3; 14 specimens of in the superior volsella; thus, they do not represent T. sp. ZUR-6; and one specimen of T. sp. ZUR-7. R. truncatocaudata. Likewise, differences in the Xestochironomus - Four species were collected, genitalia indicate that R. sp. ZUR-1 is not conspe- one undescribed. I examined the holotype of X. cific with R. manauara. ankylis, described from Guatemala by Sublette & Stempellinella - Four specimens of one unde- Sasa (1994); X. gilvus and X. latilobus were described species were collected. The genus was re- scribed from Venezuela by Borkent. Xestochirono- cently review by Ekrem (2007). mus sp. ZUR-1, represented by seven specimens, will key to couplet 6 in Borkent (1984) but is nei- Stenochironomus - Ten species of Stenochirono- ther X. subletti Borkent nor X. gilvus Borkent; it mus were collected, only one of which was posi- differs in having a very long, thin, anal point and tively identified as a previously described species: almost straight superior volsellae. S. nudipupa is known from Brazil, Costa Rica, Ecuador and Venezuela (Borkent 1984; Reis et al. CHIRCHIR -1 - This taxon is represented by 2013). A single specimen of S. cf. varius appears one damaged specimen, missing its antennae and to be S. varius but lacks any coloration on its fore all tarsomeres. The wings are bare, the squama is femur; it does have a darkened fore coxa and tro- fringed and the antepronotal lobes do not appear to chanter, and may represent a slight color variation meet medially. The gonostylus has an apical row of the species. Since Borkent’s (1984) excellent of medially directed setae, all of which are broken revision of Stenochironomus and its allies, numer- off. The specimen also lacks a subapical seta on the ous additional species have been described from superior volsella but has a subapical pit/sensillum, the Neotropics; Dantas et al. (2016) provided the from which I can observe no indication of a broken most recent key for Neotropical Stenochironomus. or lost seta. This taxon may represent a new genus Stenochironomus sp. ZUR–1 (six specimens) is or be an aberrant Stenochironomus with separated similar to S. impendens Borkent, but has a different tibial spurs (these spurs are fused in Stenochirono- abdominal color pattern; S. sp. ZUR–2 (one speci- mus); the specimen lacks the posteromedial patch men) has a dark, broad anal point, but has differ- of short setae on the gonostylus found on Xestochi- ent thoracic and abdominal color patterns from any ronomus. More, undamaged, material is needed. described species; S. sp. ZUR–3 (16 specimens) Discussion will key to S. sebastiao Andersen et al. (Andersen et al. 2008) in Dantas et al. (2016) but has a short- The Zurquí study has produced an impressive list er, thicker superior volsella and other hypopygial of chironomid taxa, the majority of which (72%) differences; S. sp. ZUR–4 (five specimens) and S. are undescribed. Noticeable in their absence were sp. ZUR–5 (one specimen) somewhat resemble some common and widespread taxa such as Ab- S. leptopus (Kieffer) but differ in coloration and labesmyia, Chironomus, Coelotanypus, Goeldi- genitalic structures; S. sp. ZUR–6 (two specimens) chironomus, Oukuriella and Tanypus. This is no is entirely pale like S. palliaculeatus Borkent, but doubt due to a dearth of suitable habitats, as many has a stouter superior volsella; S. sp. ZUR-7 (two of those genera are more common in ponds and specimens) has an dark thorax like S. varius, but lakes, especially those with a bit of eutrophication; has numerous darkened abdominal tergites; and S. no such water bodies were present at Zurquí. sp. ZUR–8 (two specimens) is unusual in having a Many of the taxa collected are considered terrestri- globose/semi-pediform superior volsella. al, semi-terrestrial or phytotelmatic; most of these Tanytarsus - Tanytarsus was by far the most belong with the subfamily Orthocladiinae: Antillo- common tanytarsine collected, with 140 speci- cladius, Bryophaenocladius, Compterosmittia, Li- mens representing nine species. The majority did purometriocnemus, Mesosmittia, Paraphaenocla- not fit descriptions of any known species, except dius, Pseudosmittia and probably Diplosmittia; at T. cf. capaira (two specimens) resembles T. ca- least one Polypedilum, P. panacu, is phytotelmatic. paira described by Trivinho-Strixino & Strixino

11 It is unfortunate that the project’s protocol did not technicians (see Borkent & Brown 2015) at INBio include collecting the immature stages of Chirono- sorted material to family; Annia Picado selected midae, but a lack of funding (only half the request- specific specimens and prepared the slides. Robin ed amount for the grant was provided) made the E. Thompson, Curator of the Insect Collection laborious process of collecting larvae and pupae, at the University of Minnesota, St. Paul, MN, and rearing them to adulthood, not possible. The made important types and other specimens avail- likelihood that many of the taxa were probably ter- able from the Sublette Collection housed there. restrial or semi-terrestrial (in wet moss, etc.), en- Broughton Caldwell reviewed an earlier draft and tailing an even more time-consuming process than helped improve this paper – many thanks, MB! Art collecting the immature stages from water, com- Borkent and Torbjørn Ekrem made many sugges- pounded the problem. One taxon collected in this tions which are incorporated in this paper. Thanks survey, a member of Parochlus araucanus group, also to Rick Jacobsen for discussions on Manoa, might have been identifiable to species if the pu- Pseudochironomus and Riethia, and to Luiz Pinho pal exuviae had been collected because the only for discussions on Diplosmittia. My wife Linda species-specific characters for this group are found helped immensely by utilizing her typing skills, far in the pupae (Brundin 1966). superior to mine, entering the majority of the iden- tifications and other data into spreadsheets. Thanks As is demonstrated in this paper, the Neotropical also to the two anonymous reviewers whose com- Chironomidae (actually, all Diptera in the Neo- ments improved the paper and indirectly led me to tropics) remain poorly known - and their immature some papers I had missed. stages, often used in environmental assessments in the Holarctic, even more so. Although Coffman This research was partially funded by US Nation- et al. (1993) reported 266 species of chironomids al Science Foundation grant DEB 1145890 to B. from 13 streams in northwestern Costa Rica, based Brown and A. Borkent. solely on pupal exuviae, none of those taxa have References been supplied with an available name. Chirono- mid taxonomy relies mostly on characters of the Andersen, T. 1996. New species of Diplosmittia male, especially the male genitalia, for species Sæther, 1981 from Costa Rica (Chironomidae, delimitation. Thus, collecting the immature stag- Orthocladiinae). - Acta Zoologica Academiae es must include rearing (or otherwise associating Scientiarum Hungaricae 42: 127–132. those stages, as with pharate pupae) to adulthood Andersen, T. & Mendes, H.F. 2002. Neotropical in order to associate those stages with the gener- and Mexican Mesosmittia Brundin, with the ally more better known adults, the bearers of the description of four new species (Insecta, Dip- species’ names. Alternatively, the use of suitable tera, Chironomidae). - Spixiana 25: 141–155. molecular markers (e.g. DNA bar codes), as shown by Trivinho-Strixino & Pepinelli (2015), may also Andersen, T., Mendes, H.F. & Pinho, L.C. 2008. help to associate life stages with available names. Two new species of Stenochironomus Kieffer, 1919 from the Brazilian Atlantic rain forest While 137 species from a four-hectare area may (Diptera: Chironomidae). - Studia Dipterolog- seem impressive, it pales with the numbers from ica 14: 263–269. some of the other families from the ZADBI project. Chironomidae only placed eleventh on the Andersen, T., Pinho, L.C. & Mendes, H.F. 2016. list of total species collected. The Cecidomyiidae Two new species of Lipurometriocnemus topped the list, with about 800 species found. A Sæther from Brazil (Diptera: Chironomidae, more complete analysis of these numbers, plus Orthocladiinae). - Revista Biotemas 29: 37-45. other aspects of diversity, biogeography, ecology, Andersen, T. & Sæther, O.A. 1996. New species etc., will be presented in a series of multi-authored and records of Beardius Reiss et Sublette (Dip- papers currently being written about the ZADBI tera: Chironomidae). - Annales de Limnologie project. 32: 33–44. Acknowledgements Andersen, T., Sæther, O.A. & Contreras-Ramos, Many thanks go to Art Borkent (Research Asso- A. 2011. New species and records of Nandeva ciate, Royal British Columbia Museum and the Wiedenbrug, Reiss et Fittkau. - Zootaxa 3136: American Museum of Natural History) and Brian 45–60. Brown (Natural History Museum of Los Angeles Borkent, A. 1984. The systematics and phylogeny County, Los Angeles, CA) for developing, organ- of the Stenochironomus complex (Xestochi- izing and supervising the ZADBI project. Several ronomus, Harrisius, and Stenochironomus)

12 (Diptera: Chironomidae). - Memoirs of the genus. - Zootaxa 3580: 43-55. Entomological Society of Canada 128: 1–269. Ekrem, T. 2007. A taxonomic revision of the ge- Borkent, A. & Brown, B.V. 2015. How to in- nus Stempellinella (Diptera: Chironomidae). - ventory tropical flies (Diptera) – One of the Journal of Natural History 41: 1367–1465. megadiverse orders of insects. - Zootaxa 3949: Epler, J.H. 1986. A novel new Neotropical Nano- 301–322. cladius (Diptera: Chironomidae), symphoretic Brown, B.V., Borkent, A., Cumming, J.M., Wood, on Traverella (Ephemeroptera: Leptophlebii- D.M., Woodley, N.E. & Zumbado, M.A. (eds.) dae). - The Florida Entomologist 69: 319–3327. 2009. Manual of Central American Diptera. Epler. J.H. 1988. Biosystematics of the genus Di- Volume 1. NRC Research Press, Ottawa, On- crotendipes Kieffer, 1913 (Diptera: Chirono- tario, Canada. 714 pp. midae: Chironominae) of the World. - Mem- Brown, B.V., Borkent, A., Cumming, J.M., Wood, oirs of the American Entomological Society D.M., Woodley, N.E. & Zumbado, M.A. (eds.) 36:1–214. 2010. Manual of Central American Diptera. Epler, J.H. 2001. Identification Manual for the Volume 2. NRC Research Press, Ottawa, On- larval Chironomidae (Diptera) of North and tario, Canada. 728 pp. South Carolina. A guide to the taxonomy of Brundin, L. 1966. Transantarctic relationships and the midges of the southeastern United States, their significance, as evidenced by chironomid including Florida. Special Publication SJ2001- midges. With a monograph of the subfamilies SP13.- North Carolina Department of Environ- Podonominae and Aphroteniinae and the aus- ment and Natural Resources, Raleigh, NC, and tral Heptagyiae. - Kungliga Svenska Vetenska- St. Johns River Water Management District, psakemiens Handlingar 11: 1–472. Palatka, FL. 526 pp. Coffman, W.P., de la Rosa, C.L., Cummins, K.W. Epler, J.H. 2010. Phytotelmatocladius, a new ge- & Wilzbach, M.A. 1993 [1992]. Species rich- nus from bromeliads in Florida and Brazil ness in some Neotropical (Costa Rica) and (Diptera: Chironomidae: Orthocladiinae). In Afrotropical (West Africa) lotic communities Ferrington, L.C., Jr. (ed.) Proceedings of the of Chironomidae (Diptera). - Netherlands Jour- XV International Symposium on Chironomi- nal of Aquatic Ecology 26: 229–237. dae. Chironomidae Research Group, Univer- sity of Minnesota, St. Paul, MN. pp. 285–293 Cranston, P.S., Martin, J., Mulder, M. & Spies, M. 2016. Clarification of Einfeldia Kieffer, 1922 Epler, J.H., Ekrem, T. & Cranston, P.S. 2013. 10. (Diptera: Chironomidae) with E. australiensis The larvae of Chironominae of the Holarctic (Freeman, 1961), comb. n. based on immature Region — Keys and diagnoses. In Andersen, stages. - Zootaxa 4158: 491–506. T., Cranston, P. S. & Epler, J. H. (Sci. eds): The larvae of Chironomidae (Diptera) of the Cranston, P.S. & Oliver, D.R. 1988. Additions Holarctic Region — Keys and diagnoses. In- and corrections to the Nearctic Orthocladiinae sect Systematics & Evolution, Supplement 66: (Diptera: Chironomidae). - Canadian Entomol- 1–571. pp. 387–556. ogist 120: 425–462. Erwin, T.L. 1982. Tropical forests: their richness Dantas, G.P.S., Hamada, N. & Mendes, H.F. 2016. in Coleoptera and other arthropod species. - Contribution to the knowledge of Stenochi- The Coleopterists Bulletin 36: 74–75. ronomus Kieffer (Diptera, Chironomidae) from Brazil: seven new species and descrip- Ferrington, L.C., Jr. & Sæther, O.A. 2011. A re- tions of females and immatures of some pre- vision of the genera Pseudosmittia Edwards, viously known species. - Zootaxa 4117: 1–47. 1932, Allocladius Kieffer, 1913, and Hydros- mittia gen. n. (Diptera: Chironomidae, Ortho- Donato, M. 2011. A new species of Bryophaeno- cladiinae). - Zootaxa 2849: 1–314. cladius (Diptera: Chironomidae) from Argen- tina. - Revista de la Sociedad Entomológica Giłka, W. & Zakrzewska, M. 2013. A contribu- Argentina 70: 207–212. tion to the systematics of Neotropical Tany- tarsus van der Wulp: first descriptions from Donato, M., Siri, A & Mauad, M. 2012. Descrip- Ecuador (Diptera: Chironomidae: Tanytarsini). tion of a new species of the genus Onconeura -Zootaxa 3619: 453–459. Andersen et Sæther (Diptera: Chironomidae) from Argentina with a cladistic analysis of the Grodhaus, G. 1987. Endochironomus Kieffer,

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15 Wiedenbrug, S. & Silva, F.L. 2016. Diplosmit- tia caribensis, a new Orthocladiinae (Diptera: Chironomidae) from the Dominican Republic. – Zootaxa 4103: 71–74.

APPENDIX 1 List of Chironomidae collected at Zurquí de Moravia, Costa Rica, 2012–2013

PODONOMINAE Cricotopus sp. ZUR-5 Parochlus araucanus Brundin, 1966, group sp. Cricotopus sp. ZUR-6 ZUR-1 Cricotopus sp. ZUR-7 Cricotopus sp. ZUR-8 TANYPODINAE Diplosmittia forficata Andersen, 1997 Procladius (P.) sp. ZUR-1 Diplosmittia sp. ZUR-1 Thienemannimyia (T.) sp. ZUR-1 Diplosmittia sp. ZUR-2 Zavrelimyia (Paramerina) fasciata (Sublette Eukiefferiella sp. ZUR-1 & Sasa, 1994) Eukiefferiella sp. ZUR-2 Eukiefferiella sp. ZUR-3 ORTHOCLADIINAE Gravatamberus guatemaltecus Mendes & Anders- Antillocladius antecalvus Sæther, 1981 en, 2008 Antillocladius arcuatus Sæther, 1982 Irisobrillia longicosta Oliver, 1985 Antillocladius pluspilalus Sæther, 1981 Krenosmittia sp. ZUR-1 Bryophaenocladius cf. psilacrus Sæther, 1982 Limnophyes guatemalensis Sublette & Sasa, 1994 Bryophaenocladius sp. ZUR-1 Limnophyes mariae Sublette & Sasa, 1994 Bryophaenocladius sp. ZUR-2 Lipurometriocnemus glabalus Sæther, 1981 Bryophaenocladius sp. ZUR-3 Lipurometriocnemus sp. ZUR-1 Bryophaenocladius sp. ZUR-4 Lipurometriocnemus sp. ZUR-2 Compterosmittia nerius (Curran, 1930) Lipurometriocnemus sp. ZUR-3 Compterosmittia sp. ZUR-1 Litocladius chavarriai Mendes, Andersen & Hagenlund, 2011 Compterosmittia sp. ZUR-2 Lopescladius (Cordiella) sp. ZUR-1 Corynoneura ferelobata Sublette & Sasa, 1994 Lopescladius (Cordiella) sp. ZUR-2 Corynoneura guanacaste Wiedenbrug, Lamas & Trivinho-Strixino, 2012 Mesosmittia patrihortae Sæther, 1986 Corynoneura sp. ZUR-1 Mesosmittia sp. ZUR-1 Corynoneura sp. ZUR-2 Metriocnemus costatus Sublette & Sasa, 1994 Cricotopus sp. ZUR-1 Metriocnemus sp. ZUR-1 Cricotopus sp. ZUR-2 Metriocnemus sp. ZUR-3 Cricotopus sp. ZUR-3 Nanocladius sp. ZUR-1 Cricotopus sp. ZUR-4 Onconeura sp. ZUR-1

16 Parakiefferiella sp. ZUR-1 Einfeldia sp. ZUR-1 Parametriocnemus sp. ZUR-1 Endotribelos albatum Sublette & Sasa, 1994 Parametriocnemus sp. ZUR-2 Endotribelos grodhausi Sublette & Sasa, 1994 Parametriocnemus sp. ZUR-3 Endotribelos sp. ZUR-1 Parametriocnemus sp. ZUR-4 Nandeva latiloba Sæther & Roque, 2004 Paraphaenocladius exagitans longipes Sæther & Nilothauma sp. ZUR-1 Wang, 1995 Nilothauma sp. ZUR-2 Pseudorthocladius sp. ZUR-1 Parachironomus sp. ZUR-1 Pseudosmittia forcipata (Goetghebuer, 1921) Phaenopsectra sp. ZUR-1 Pseudosmittia windwardensis (Sæther, 1981) Polypedilum (Pentapedilum) sp. ZUR-1 Pseudosmittia sp. ZUR-1 Polypedilum (Polypedilum) panacu Mendes, An- Stictocladius sp. ZUR-1 dersen & Jocqué, 2011 Synorthocladius sp. ZUR-1 Polypedilum (Polypedilum) sp. ZUR-1 CHIRORTH-1 sp. ZUR-1 Polypedilum (Polypedilum) sp. ZUR-2 CHIRORTH-2 sp. ZUR-1 Polypedilum (Polypedilum) sp. ZUR-3 CHIRORTH-3 sp. ZUR-1 Polypedilum (Tripodura) apicatum Townes, 1945 CHIRORTH-4 sp. ZUR-1 Polypedilum (Tripodura) luteopedis Sublette & Sasa, 1994 CHIRORTH-4 sp. ZUR-2 Polypedilum (Tripodura) obelos Sublette & Sasa, CHIRORTH-5 sp. ZUR-1 1994 CHIRORTH-6 sp. ZUR-1 Polypedilum (Tripodura) sp. ZUR-2 CHIRORTH-7 sp. ZUR-1 Polypedilum (Uresipedilum) sp. ZUR-1 CHIRORTH-8 sp. ZUR-1 Rheotanytarsus cf. contrerasi Andersen & Sæther, CHIRORTH-9 sp. ZUR-1 2000 CHIRORTH-10 sp. ZUR-1 Rheotanytarsus cf. guanacastensis Kyerematen & CHIRORTH-11 sp. ZUR-1 Andersen, 2002 CHIRORTH-12 sp. ZUR-1 Rheotanytarsus scutulatus Kyerematen & Anders- en, 2002 CHIRORTH-13 sp. ZUR-1 Rheotanytarsus subtilis Kyerematen & Andersen, CHIRORTH-14 sp. ZUR-1 2002 CHIRORTH-15 sp. ZUR-1 Rheotanytarsus sp. ZUR-1 CHIRORTH-16 sp. ZUR-1 Riethia sp. ZUR-1 Stempellinella sp. ZUR-1 CHIRONOMINAE Stenochironomus nudipupa Borkent, 1984 Beardius triangulatus Andersen & Sæther, 1996 Stenochironomus cf. varius Borkent, 1984 Beardius sp. ZUR-1 Stenochironomus sp. ZUR-1 Caladomyia sp. ZUR-1 Stenochironomus sp. ZUR-2 Caladomyia sp. ZUR-2 Stenochironomus sp. ZUR-3 Cryptochironomus sp. ZUR-1 Stenochironomus sp. ZUR-4 Dicrotendipes sp. ZUR-1 Stenochironomus sp. ZUR-5

17 Stenochironomus sp. ZUR-6 Tanytarsus sp. ZUR-5 Stenochironomus sp. ZUR-7 Tanytarsus sp. ZUR-6 Stenochironomus sp. ZUR-8 Tanytarsus sp. ZUR-7 Tanytarsus cf. caipira Trivinho-Strixino & Strix- Tanytarsus sp. ZUR-8 ino, 2007 Xestochironomus ankylis Sublette & Sasa, 1994 Tanytarsus sp. ZUR-1 Xestochironomus gilvus Borkent, 1984 Tanytarsus sp. ZUR-2 Xestochironomus latilobus Borkent, 1984 Tanytarsus sp. ZUR-3 Xestochironomus sp. ZUR-1 Tanytarsus sp. ZUR-4 CHIRCHIR-1 sp. ZUR-1

Article submitted 19. February 2017, accepted by Torbjørn Ekrem 7. April 2017, published 21. April 2017.