Spies Etal 2009

CHIRONOMIDAE (NON-bItINg MIDgEs)
Martin Spies, Trond Andersen, John H. Epler, and Charles N. Watson, Jr.
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Fig. 30.1. Male of Chironomus plumosus (Linnaeus), (Holarctic, MND, fig. 29.1).
Diagnosis
Adults small to medium-sized, about 113 mm long. Morphology diverse, some (especially marine) taxa with modifications or reductions to structures of the head, wings, legs, or terminalia. Ocelli absent; frontal tubercles present (e.g., Fig. 14) or absent. Compound eyes large, often with dorsomedial extensions (e.g., Fig. 14), but never contiguous. Antennae most commonly strongly sexually dimorphic; scape a flat ring; pedicel globose (especially enlarged in male); males with 615 flagellomeres, females with 47 or 1014; male antennae usually with first and either last or penultimate flagellomere longer than others, and with flagellum densely plumose (Fig. 1), but exceptions occur in several genera. Mouthparts not piercing; female mandibles rarely present (no taxa with mandibles in Central America); proboscis rarely elongate; palpus usually elongate, with five (= four free) segments (Figs. 1214), in various taxa reduced 437
in length and/or segment number. Wing (Figs. 211) usually narrow (but see Fig. 5), broader in female (absent in some marine females); wings in resting position held to posterior, either flat (may be overlapping) or like sloping roof; costa reaching at most to tip of wing; wing margin usually reached by two to three branches of R, one of M, and two of Cu (but see Table 1 below). Legs (Figs. 3049) usually elongate with forelegs longest, midlegs shortest; first tarsomeres longer than second; males at rest often hold forelegs aloft like feelers (Fig. 1). Postnotum usually with medial longitudinal groove (Fig. 19). Abdomen usually much longer than wide, especially slender in males. Larvae (Figs. 102106) elongate, cylindrical (thoracic segments inflated in prepupae); with sclerotised, prognathous, usually non-retractable head capsule, three thoracic and nine abdominal segments, and usually anterior and posterior pair of prolegs bearing spines or claws; antennae various (long and retractable in Tanypodinae, Fig. 105); mouthparts di-
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MANUAL OF CENtRAL AMERICAN DIPtERA VOLUME 1
Table 1. Selected morphological terms differing between the present manual (after Oliver, 1981) and current chironomid terminology (after Sther, 1980). Present manual Wing crossvein r-m vein M crossvein bm-cu cubital fork vein CuA1 vein CuA2 calypter Sther (1980) RM M + M1+2 MCu FCu M3+4 Cu1 squama Present manual Male terminalia epandrium gonocoxal apodeme paramere dorsomedial lobe appendage 1 appendage 1a appendage 2 appendage 2a gonostylar tooth Sther (1980) Hypopygium tergite IX coxapodeme + sternapodeme phallapodeme + aedeagal lobe median volsella (in part) superior volsella (in part) digitus inferior volsella (in part) median volsella (in part) megaseta
rected anteriorly (prognathous), with mandibles operating in oblique to horizontal plane, and ventromedially with either transverse, usually toothed plate (hypostoma, Fig. 106; = mentum) or movable, toothed ligula (in larvae of Tanypodinae, Fig. 105); open spiracles usually absent (absent in all Central American species); terminal segments usually with paired procerci and four anal papillae (Fig. 102), sometimes also with short lateral and/or longer ventral tubuli (Fig. 103). Reductions occur, especially of prolegs and terminal appendages, for example in some non-aquatic larvae (e.g., Fig. 104). Pupae (Figs. 99101) comma-shaped or relatively straight in lateral view, abdomen much narrower than cephalothorax, more or less flattened dorsoventrally; thorax anterolaterally often with pair of thoracic horns (sac-, tube-, trumpet- or tuftlike; simple, bilobed or with few to dozens of branches); antennal sheaths elongate, curving laterally over eyes; leg sheaths separate, usually at least tips of hind leg sheaths recurved under wing pads; tip of abdomen with or without apical pair of flattened anal lobes, the latter often with lateral fringe or distal setae or spines, not stiff and bare or with midrib. Various reductions from above occur, especially in non-aquatic pupae. Adults are most often confused with culicids, chaoborids or ceratopogonids. In the former two families, the wing margin is reached by 10 vein branches (at most 7 in Chironomidae), and the costa continues around the wing. Ceratopogonidae differ from all chironomids in the combination of postnotum lacking a longitudinal groove and wing without bm-cu.
Biology
Armitage et al. (1995) have provided a comprehensive overview of chironomid biology and ecology, and a guide to much of the large and continuously growing body of literature on the various themes in which members of this insect family play a role, including those outlined below. Adult Chironomidae are non-biting; if they feed during their short lifespans of just hours to a few weeks, then only
on flower nectar or honeydew. None are known to attack or transmit diseases to humans or other living organisms, although some of their larvae live as predators, invertebrate or fish parasites, or mine in and damage plants. However, the family is so ubiquitous, successful, and productive in and around nearly all kinds of inland water bodies, in wet soil or similar accumulations of organic matter, that mass populations of Chironomidae often occur, which may interfere with human activities near such habitats or even with peoples well-being. Some species carry molecules in their hemolymph that serve to store and transport dissolved oxygen much like the hemoglobins in the blood of higher animals, enabling the larvae of such species to survive and develop in stagnant or even fouling waters. These chironomid hemoglobins rank among the most potent human allergens known, as was discovered when midge larvae (bloodworms) began to be cultivated commercially as food for fishes in human culture, and workers came into prolonged contact with substances from the dried and ground-up larval bodies. Although these hemoglobins are partially broken down and no longer physiologically active in adults of such chironomid species, their potency as allergens is little diminished. Consequently, a number of cases are on record from around the world though not yet from the Neotropical Regionof people developing mild to serious allergies due to contact with clouds of emerging or swarming adult Chironomidae. Even if no human health problems arise, midge population explosions can cause a variety of problems, from infestations of water supply systems, sewage treatment plants, or rice plantations through interference with industrial production (e.g., when swarms of chironomid adults settle on objects freshly sprayed or wet-painted) to the daily and nightly nuisance felt by residents of lake- or riverfront properties. The major factors governing chironomid productivity in a given habitat are the suitability of temperature and other environmental parameters, the food base, and the presence of competitors and/or predators. Although many species are generalists able to grow and reproduce under a relatively broad range of combinations of the above factors, a large and diverse array of more or less specialized niches has
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439 live more or less planktonically and to disperse before settling on suitable substrates with which the later instars then associate more sedentarily. This change in the way of living can be reflected in structural modifications, e.g., to the shapes of head capsule and mouthparts. Depending on the degree of specialization of a species, the second to fourth instar larvae may live on or in different or specific substrates such as muddy to rocky sediments, algal mats, the surfaces or internal tissues of rooted or floating aquatic macrophytes, immersed dead wood, or in phoretic or even ecto- to endoparasitic associations with other animals. Most larvae are microphagous, ingesting detritus, small algae or decaying plant and animal tissues. Other larvae, especially (but not exclusively) in Tanypodinae, are more or less carnivorous. Most feed opportunistically on whatever is available, but in some taxa food items are filtered or trapped with the use of selfspun strands of sticky silk (e.g., Rheotanytarsus). The larvae of many species use such silk to line parts of their sediment burrows or construct tubelike shelters, usually incorporating substrate particles such as mud, sand grains, shell, or plant fragments. Tubes are often affixed to the substrate, including on the bodies of other living and moving animals, and later serve to shelter the pupa as well. In a few genera of Orthocladiinae and Chironominae transportable cases are built, similar to those of some Trichoptera. The larvae of certain species (e.g., many Tanypodinae) are free living and do not construct tubes for themselves or their pupae. Transformation of the body from larval to pupal and adult structures begins in the fourth instar, and larvae nearing pupation are usually easily distinguishable from those in earlier stages of development by their conspicuously inflated anterior body segments. In some subfamilies (e.g., Tanypodinae) pupae are free living, active swimmers, but in most species they remain within the shelter built by the larva. Duration of the pupal stage ranges from hours to several days, after which aquatic pupae rise to the water surface for adult eclosion. Adults emerge quickly from the pupal skin (exuviae), taking at most a few minutes to unfold and stiffen their wings, although after flying off it may take a day for their cuticle to fully harden and coloration to develop its final pattern. Adult behavior varies among taxa, resulting in different patterns of resting and activities such as locomotion and mating. In most species the males form aerial swarms, often cued to specific diel periods (e.g., sunset) and/or positional markers (e.g., next to a low bush at a waterline, or over a prominent tree top). Attracted females enter the swarm and copulation ensues, either quickly in flight or for more extended periods on the ground. Mating without swarming, on a firm substrate or the water surface, is also common, the latter, for example, in many marine species. Adult morphology often reflects the mating behavior, for example in relative size and development of the male antennae and terminalia. Several different taxon-specific mating positions occur within the Chironomidae, for some of which the male terminalia are rotated around the longitudinal body axis.
also been conquered and cultivated. For example, Chironomidae are often among the few insects surviving in marine intertidal zones, in near-coastal habitats experiencing salinity fluctuations, or in waters impacted by sewage or other pollution. This versatility and adaptability of the chironomid bauplan has earned the family one of the most dominant ranks (often clearly No. 1), by numbers of individuals or species, among comparable taxa in many aquatic invertebrate communities. With this large number and high ecological diversity of species the Chironomidae also assume key roles in ecosystems and food webs, e.g., by forming large parts of the diet of other invertebrates, fish or waterfowl, by aiding the self-cleaning of habitats loaded with organic matter, or by influencing material retention/mobilization processes between sediments and the water and between aquatic and terrestrial systems. Life cycles of Chironomidae range from a few days (Nolte, 1995) to several years. This may include periods of dormancy, hibernation, or aestivation, depending on the interaction between requirements and adaptations of the animals and the conditions and cues provided by the environment. Some species inhabiting regions with arid periods can tolerate desiccation in the larval stage (Cranston & Nolte, 1996). One African Polypedilum has been shown to continue and complete normal development when rehydrated after years in the dried-out state. In tropical or subtropical and near-optimal growth conditions, development and emergence may be continuous and little synchronized, which may result in many generations per year (about 50 for a small Corynoneura in a subtropical North American stream; Benke et al., 1984). Larger species or those growing in cooler or otherwise more limiting environments often complete only one or two generations per year, with synchronized emergence periods each limited to several weeks, days or even only hours. However, few data are available on actual phenologies of Central American chironomid populations. Chironomid females deposit their eggs on various resident or floating substrates such as stones, plant stems or leaves, wood, leaf litter, or just the water surface. Eggs vary in shape and may be laid individually or in loosely to elaborately arranged and packaged masses; these characteristics are often diagnostic for particular taxa (Nolte, 1993). Egg numbers laid per female range from a few to several thousand. Egg development times and the occurrence and duration of dormancy or diapause phases in this stage are also highly variable. The larva is usually the stage in which a chironomid spends most of its lifetime. Each larva moults three times, i.e. passes through four instars, prior to pupation. Larvae may be found in habitats ranging from all kinds, sizes, and depths of open water through spring seeps and hygropetric sites, or permanently or temporarily wetted plant stands, to saturated or damp soil and similar accumulations of organic matter, such as leaf litter, the dung of larger animals, etc. In many aquatic species, the first instars are considered to
440 A number of chironomid species are parthenogenetic, either facultatively (under conditions occurring temporarily or in part of the distributional range) or obligatorily. Among the genera occurring or likely present in Central America, the following have some members showing parthenogenesis (at least in other world regions): ChironominaeMicropsectra, Paratanytarsus, Pontomyia, Tanytarsus; Orthocladiinae Bryophaenocladius, Limnophyes, Pseudosmittia, Smittia.
Classification
While monophyly of the Chironomidae appears certain (e.g., Cranston, 1995), interpretations of their relations to other families vary among different studies (Sther, 2000a). Most often, the Ceratopogonidae and/or Simuliidae are considered the most closely related. Within the Chironomidae, phylogeny and even parts of the classification are also still subject to reinterpretations (e.g., Cranston, 1995; Sther, 2000b) as additional data are continuously being gained from newly discovered taxa, life stages or character sets (e.g., gene sequences). Although chironomid systematics has incorporated information not only on the adults but also on the immature stages far more than in most other dipteran families, there is also a negative side to this: classifications and phylogenies based on different life stages often turn out to be incongruent. The lack of complete representation of all life stages in the material for many taxa, and of comprehensive revisions of all stages, is another obstacle to a truly consistent system. Eleven subfamilies are currently recognized (Sther, 2000b; Spies, 2005), of which 10 have been recorded from the Neotropical Region (Spies & Reiss, 1996), and 6 from Central America. The vast majority of speciesin the Neotropical Region about 80%, in Central America over 95% are assigned to one of the three subfamilies Chironominae, Orthocladiinae and Tanypodinae. Most of the remaining subfamilies contain one or a few species in one or several genera (Central America: Buchonomyiinae, Telmatogetoninae), but some do attain considerable richness in certain world regions (e.g., Podonominae in relatively cool, montane areas especially of the southern Neotropical Region and New Zealand; Diamesinae in North America). In the Chironominae, three tribes are recognized (Chironomini, Pseudochironomini, Tanytarsini). All have Central American representatives, which are distributed among 49 genera currently recorded from the region. The Tanypodinae are divided into six tribes (Coelotanypodini, Macropelopiini, Natarsiini, Pentaneurini, Procladiini, Tanypodini), but comparatively few genera (Central America: all tribes except Natarsiini, 17 genera). In the Orthocladiinae there are no unanimously accepted tribes, but large and growing numbers of genera (28 in Central America). About 5000 species of Chironomidae worldwide are currently considered scientifically valid, but estimates for the total number of extant species range from about 8000 to 20 000 (Coffman, 1995). Many large regions known or ex-
pected to support high chironomid diversity are still poorly studied in comparison to those in the Holarctic Region. For example, about 1100 named species are recognized from North America (Oliver et al., 1990; Oliver & Dillon, 1994), but only about 800 from the Neotropical Region (Spies & Reiss, 1996; database maintained by M. Spies), even though the latter includes Amazonia and similar areas with many known but still undescribed species (e.g., vast material in the collection of the Zoologische Staatssammlung Muenchen, ZSM, gathered by E.J. Fittkau and collaborators). The approximately 200 species so far recorded from Central America certainly are a gross underestimation of the actual fauna as well, considering the geographic extent and physiographic diversity of the region. Many species have been collected and recognised as newincluding much material in the collections accessible to the present authorsbut are still undescribed due to shortages of manpower and time. Some require the formation of new genera, but the latter could not yet be treated in the present manual. A total chironomid richness close to 1000 species in Central America would not be surprising.
Identification
An up-to-date, comprehensive guide to the literature on taxonomy and identification of the Chironomidae is not available. Cranston (1995) offers separate keys for adult males, females, pupae, and larvae to the subfamilies of Chironomidae (except for Usambaromyiinae Andersen & Sther, 1994, which contains a single genus and species so far known only from Africa). Many Central American larvae should key reasonably well in Epler (2001; see also http://home.comcast.net/~johnepler3/index.html) or Coffman & Ferrington (1996), if one stays mindful of the differences between regions in faunal composition even at the generic level. Wiederholm (1983) also contains much useful information, but is partially outdated. For pupae and pupal exuviae, Coffman & Ferrington (1996) or Wiederholm (1986) may be tried, with caution because the geographic scopes and centers of origin of these works are even farther away from Central America than Eplers. Sther (1977b) gave a large-scale overview of chironomid female terminalia morphology. This includes keys and diagnoses for genera, but due to their age and inadvertent limitations on material studied these do not always lead to reliable identifications, especially with Neotropical specimens. Monographs exist on adults of certain chironomid subfamilies or tribes (Townes, 1945 for Nearctic Chironomini; Fittkau, 1962 for several Tanypodinae tribes; Roback, 1971 for Nearctic Tanypodinae), and some of the species treated there might also be found in Central America. On the other hand, some of the contents of these works are outdated, and the different geographic scopes must be kept in mind as well. For additional monographs on individual genera see the references provided in the Synopsis section below. Spies & Reiss (1996) cataloged information and references for all
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441 be difficult unless a certain life stage is known (usually the adult male or the pupa). Reliable associations obtained by individual rearings multiply the chances for identification, and will improve our knowledge of species, their ecology and diversity much more quickly than the usual grab-andpreserve sampling methods, which yield such associations too rarely and only by chance. Under subtropical conditions with relatively high natural water temperatures, rearing often requires little more effort from the researcher than to place one live prepupal larva or pupa into a vial with a little water from the source, monitor the animals development, wait for the adult to emerge and fully harden, then preserve it along with the cast skins it left in the vials water. Spies (2000b) has shown that this approach may be necessary but on the other hand sufficient to identify nearly all important components of a chironomid fauna even where notoriously difficult genera (e.g., Chironomus, Cricotopus) are present with several species each. Without such life-stage associations, genus identifications of isolated immature stages and some adults must often be considered as tentative. Consequently for Central America, a key can be offered here for adult male specimens only, because for a significant share of the species other life stages are not or insufficiently known. The identification of adult femalesfrequently collected and described with the malesoften depends on minute details of external and internal genitalia, which for many taxa have not yet been elaborated. Reliable associations of immature stages have been established for even fewer species. In most cases, identification will require complete or partial slide-mounting of the specimen, usually involving dissection and the removal of internal tissuese.g., from the larval head capsule, the adult thorax and terminaliaby maceration in 510% KOH. Detailed descriptions of techniques can be found in Epler (2001), along with much additional practical information and advice on rearing, steps to increase and assure the quality of ones results, etc. See also the Preservation and mounting section in Wiederholm (1989). Chironomid researchers worldwide, although not formally organized in an association, have for decades formed a close community, with international symposia held since 1964 (presently every three years). There is The Chironomid Home Page (http://insects.ummz.lsa.umich. edu/~ethanbr/chiro/), free for anybody to access or contribute, and the Chironomus Newsletter on Chironomidae Research (published once a year as a PDF file, downloadable from the Home Page), which includes annual bibliographies. Additional websites present information on subsections of the chironomid realm, defined either geographically or by special topics (e.g., subfossil midge remains); links to these can also be found on The Chironomid Home Page.
taxa known from the entire Neotropical Region at that time, including data on the life stages treated in each original species description and distribution records by country. However, many additional taxa have since been published from the Neotropical Region: for example, the total number of named species has increased by 15% in less than 10 years. On the scale of countries, the following faunal overviews have been published: Mexico (Contreras Ramos & Andersen, 1999; Andersen et al., 2000); Guatemala: Sublette & Sasa (1994); Costa Rica: Watson & Heyn (1993). Up-to-date lists can be produced from the database maintained by the first author upon request. The abundance and diversity of Chironomidae demonstrate the importance of the family in the biota, but they are also the reason why in many ecological studies the information content is under-utilized. Often, the lack of descriptions and keys to a local fauna precludes species determinations, or workers choose to ignore the Chironomidae in favor of groups (e.g., Ephemeroptera, Plecoptera, Trichoptera) that are more limited in number and diversity and thus perhaps less informative, but whose members are supposedly more easily identified. On the other hand, if species identifications are possible, chironomids make a highly valuable tool for the surveying and monitoring of both pristine and impacted habitats. One of the easiest, most effective and least destructive techniques for such studies is the interception of current or skimming of water surfaces for cast pupal skins (exuviae) and emerging or drowned adults. This method will become even more valuable in Central America as more pupae are associated with the respective male adults. The latter is still the stage to which most scientific species names of Chironomidae are tied by virtue of the original descriptions. Immature stages of aquatic species are taken with all kinds of standard invertebrate samplers, in bulk sediment or vegetation grabs, orespecially from small or special habitatsby hand with forceps, pipettes, etc. Adults are readily collected with aerial or sweep nets, in light or Malaise traps, or during emergence, with eclectors for species with terrestrial larvae or with cone-shaped traps the wide ends of which are submerged below a water surface (the latter method may also yield pupal exuviae). The usual preservative is 7080% alcohol. To avoid problems such as breakage of limbs, vials should not be overfilled, and should be stoppered such that air bubbles are kept away from the specimens. Where special color patterns are to be documented representative animals should be photographed prior to immersion, or adults carefully dried and pinned. In Central America and elsewhere, where the chironomid fauna will likely remain poorly known for some time, the best way to study and identify these animals is to rear larvae through to the adult stage individually. Some species can be impossible to identify, and even assignment to a genus can
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Key to the adult males of the genera of Chironomidae of Central America

Genera with names placed between square brackets in the key have not been recorded from Central America, but are considered likely to occur there by the present authors. Since the publication of Sther (1980), most discussions of morphological structures in Chironomidae have used the terms preferred in that glossary (older terms are explained there as well). This current chironomid terminology differs 1. 2. 3. 4. from the one mandated for the present manual in many critical elements, notably concerning certain wing veins, adult thoracic sclerites, parts of the male terminalia, and structures of the larval head capsule. In order to facilitate the readers use of the recent chironomid literature, Table 1 presents both sets of terms for those used in the key.
Wing with crossvein bm-cu present (Figs. 24). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Crossvein bm-cu absent (Figs. 511) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Wing vein R2+3 absent (Figs. 3, 4); either with costa reaching wing tip (Fig. 3), or with terminalia including digitiform gonocoxite appendage 2 and complex aedeagus (Fig. 50) . . . . . . . . . . . . . . 3 R2+3 usually present (Fig. 2), if absent, then costa ending distinctly proximal of wing tip and terminalia not as above; TANYPODINAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Crossvein bm-cu close to middle of wing (Fig. 3); first axillary sclerite (plate) of wing without setae; PODONOMINAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Parochlus Enderlein Crossvein bm-cu closer to wing base (Fig. 4); first axillary sclerite with setae; BUCHONOMYIINAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Buchonomyia Fittkau Wing vein R2+3 absent; all tarsomeres 4 cordiform (Fig. 30); terminalia with conspicuous aedeagus projecting beyond epandrium and flanked by pair of sclerotized plates; TELMATOGETONINAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 R2+3 usually present (Figs. 711), if absent and tarsomeres not elongate, cylindrical, then terminalia without conspicuous, projecting aedeagus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Tarsomeres 5 simple; palpus with five (= four free) segments (similar to Figs. 1214); thorax with numerous acrostichal setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Thalassomya Schiner Tarsomeres 5 deeply trilobed (Fig. 30); palpus with no more than three (two free) segments; thorax without acrostichal setae (Fig. 20) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Telmatogeton Schiner Gonostylus usually rigidly fused to gonocoxite (Figs. 6082), and foretarsomere 1 longer than or subequal to foretibia (Fig. 32); if gonostylus somewhat movable against gonocoxite or foretarsomere 1 slightly shorter than foretibia, then hind tibial combs consisting of basally fused spines (similar to Figs. 4547); CHIRONOMINAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31 Gonostylus flexibly attached to gonocoxite (Figs. 8398); foretarsomere 1 shorter than foretibia (Fig. 34); hind tibial comb, when present, consisting of separate spiniform setae (Figs. 33, 42, 43); ORTHOCLADIINAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102 All tarsi with fourth tarsomere cordiform (similar to Fig. 31); hind tibia with comb setae in two parallel rows; wing vein R2 not connected to R3; COELOTANYPODINI . . . . . . . . . . . . . . . . . . 8 Fourth tarsomeres cylindrical; hind tibial comb in a single row (Figs. 3941) or absent; R2 connected to R3 (Fig. 2) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Scutum with small, mediocentral tubercle; wing with cubital fork close to crossvein bm-cu. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Coelotanypus Kieffer Scutal tubercle absent; distance from bm-cu to cubital fork at least as long as one-quarter of CuA2 (similar to Fig. 2) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
5. 6.
7. 8.
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9. 10.
Thoracic postnotum with dorsal setae; eye with well-developed dorsomedial extension (similar to Fig. 14) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Clinotanypus Kieffer Postnotum without dorsal setae; eye extension absent. . . . . . . . . . . . . . . . . Naelotanypus Roback Wing with cubital fork distinctly distal to crossvein bm-cu (Fig. 2) . . . . . . . . . . . . . . . . . . . . . . 11 Cubital fork adjacent or proximal to bm-cu . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
Figs. 30.211. Wings: dorsal view of (2) Procladius freemani Sublette, (Nearctic, MND, fig. 29.2); (3) Parochlus kiefferi (Garrett), (Holarctic, MND, fig. 29.6); (4) Buchonomyia brundini Andersen & Sther, (Andersen & Sther, 1995, fig. 1D); (5) Clunio sp., (Nearctic, MND, fig. 29.10); (6) Corynoneura sp., (Nearctic, MND, fig. 29.11); (7) Cardiocladius obscurus (Johannsen), (Nearctic, MND, fig. 29.12); (8) Smittia sp., (Nearctic, MND, fig. 29.14); (9) Chironomus riparius (Meigen), (Holarctic, MND, fig. 29.16); (10) Micropsectra sp., (Nearctic, MND, fig. 29.17); and (11) Stempellina sp., (Nearctic, MND, fig. 29.18). Abbreviations: calyp, calypter; Cu fk, cubital fork.
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Figs. 30.1221. Heads, antenna, and thoraces: frontal views of heads (antennal flagella omitted) of (12) Nanocladius sp., (Nearctic, MND, fig. 29.20); (13) Orthocladius consobrinus (Holmgren), (Holarctic, MND, fig. 29.21); and (14) Glyptotendipes barbipes (Staeger), (Holarctic, MND, fig. 29.22); apex of antenna of (15) Smittia sp., (Nearctic, MND, fig. 29.23); dorsal views of antepronotum of (16) Chironomus sp., (Nearctic, MND, fig. 29.35); (17) Axarus scopula (Townes), (Nearctic, MND, fig. 29.36); and (18) Glyptotendipes barbipes (Staeger), (Holarctic, MND, fig. 29.37); dorsal view of thorax of (19) Ablabesmyia sp., (Nearctic, MND, fig. 29.26); lateral views of thorax of (20) Telmatogeton alaskensis Coquillett, (MND, fig. 29.39); and (21) Limnophyes brachytomus (Kieffer), (MND, fig. 29.40). Abbreviations: aanepst, anteanepisternum; acr s, acrostichal setae; anapl sut, anapleural suture; aprn lb, antepronotal lobe; cx, coxa; dc s, dorsocentral setae; epm, epimeron; epm s, epimeral setae; kepst, katepisternum; manepst, medioanepisternum; lng gr, longitudinal groove; panepst, postanepisternum; pn, postnotum; pocl s, postocular setae; sct, scutum; sctl, scutellum; sctl s, scutellar setae; spal s, supraalar setae; vt s, vertical setae.
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445
11. 12.
Distance from bm-cu to cubital fork at most as long as one-third of CuA2; wing usually spotted; scutal tubercle present; TANYPODINI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Tanypus Meigen Distance from bm-cu to cubital fork at least as long as one-half of CuA2 (Fig. 2); wing clear or banded; scutal tubercle present or absent; PROCLADIINI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Gonostylus strongly hooked, usually with one or two ventral (inner) branches or lobes (Fig. 51); thorax, legs, and abdomen with striking patterns of contrasting pigmentation; eyes iridescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Djalmabatista Fittkau Gonostylus not strongly hooked, at most with one ventral lobe (Figs. 53, 54); coloration less conspicuous at least on thorax and legs, although scutal stripes (vittae) and apices of femora and tibiae may be darkened; eyes not iridescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 Scutal tubercle present . . . . . . . . . . . . . . Laurotanypus Oliveira, Messias & da Silva-Vasconcelos Scutal tubercle absent; Procladius Skuse. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Gonostylus with posterior (outer) heel, ventral (inner) lobe absent (Fig. 54) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Procladius (Holotanypus) Roback Gonostylus without heel, ventral lobe present or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 Gonostylus without ventral lobe . . . . . . . . . . . . . . . . . . . . . . . . . . Procladius (Procladius) Skuse Gonostylus with ventral lobe (Fig. 53) . . . . . . . . . . . . . . . . . . . Procladius (Psilotanypus) Kieffer Wing with costa produced beyond R4+5 for a distance usually at least as long as crossvein r-m; postnotal setae present or absent; MACROPELOPIINI1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 Costa ending at R4+5, or at most produced for a distance shorter than r-m; postnotal setae absent; PENTANEURINI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 Scutal tubercle and postnotal setae present; palpal segment 3 semiglobose, with strong setae on inner margin; foretibia without comb; epandrium without setae . . . . . . Fittkauimyia Karunakaran Scutal tubercle absent, postnotal setae present; palpal segment 3 cylindrical, with normal setae on inner margin; foretibia without comb; epandrium with setae; South America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Guassutanypus Roque & Trivinho-Strixino] Scutal tubercle and postnotal setae absent; palpal segment 3 cylindrical, with normal setae on inner margin; foretibia with comb; epandrium with setae . . . . . . . . . . . . . . . . . Alotanypus Roback Each tibia with at least three dark bands; wing with dark spots; rows of acrostichal setae diverge around prescutellar area (Fig. 19); terminalia with complex dorsomedial lobe carrying chitinized rods and setiferous lobes; gonostylar tooth spoon shaped, arising subapically (Fig. 57) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Ablabesmyia Johannsen Tibiae at most with single band near one end; if rows of acrostichals divergent, then only around anterior part of prescutellar area; if terminalia with dorsomedial lobe, then without chitinized rods; gonostylar tooth tapering to point, arising apically. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
13. 14. 15. 16. 17.
18.
The Neotropical Macropelopiini are not well known. Genus concepts and diagnoses in the tribe are based on combinations of adult characters occurring in Holarctic species, and also make extensive use of larval and pupal features (Fittkau, 1962; Roback, 1971, 1978; Wiederholm, 1989). Most Neotropical Macropelopiini species have been described from adult specimens only, and many do not fit any recognized genus (see the list of Unplaced valid species in Tanypodinae in Spies & Reiss 1996). In Central America, one such unassignable species is Macropelopia roblesi Vargas, 1946 from Mexico. Moreover, the discovery of previously unknown immature stages can call into question generic placements made earlier on the basis of adult characters alone (Watson, 1999). Currently, two genera can be recognized in Central America, but undescribed species are known (and more are likely to be found) that cannot be placed in existing genera and will not fit the present key. There is a real need for rearing to obtain specific associations of larvae, pupae and adults in order to understand the Neotropical Macropelopiini.
446
Figs. 30.2234. Thoraces: lateral views of pronotum and scutum of (22) Conchapelopia pallens (Coquillett), (Nearctic, MND, fig. 29.41); (23) Orthocladius consobrinus (Holmgren), (Holarctic, MND, fig. 29.42); (24) Nanocladius sp., (Nearctic, MND, fig. 29.44); (25) Parakiefferiella sp., (Nearctic, MND, fig. 29.45); (26) Antillocladius sp., (illustrated by T. Andersen); (27) Omisus pica Townes, (Nearctic, Townes, 1945, fig. 228); (28) Stictochironomus sp., (Nearctic, MND, fig. 29.47); and (29) Polypedilum simulans Townes, (Nearctic, MND, fig. 29.48); ventral views of third to fifth hind tarsomeres of (30) Telmatogeton alaskensis Coquillett, (MND, fig. 29.49); and (31) Cardiocladius obscurus (Johannsen), (Nearctic, MND, fig. 29.50); lateral view of right foreleg of (32) Microtendipes pedellus (De Geer), (Holarctic, MND, fig. 29.52); posterior apex of hind tibia of (33) Corynoneura sp. (Nearctic, MND, fig. 29.53); lateral view of right foreleg of (34) Corynoneura sp., (Nearctic, MND, fig. 29.54; arrow indicates dorsal keel on trochanter). Abbreviations: acr s, acrostichal setae; dc s, dorsocentral setae; sct tub, scutal tubercle.
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19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30.
Terminalia with dorsomedial lobe (Figs. 55, 56); Thienemannimyia group2 . . . . . . . . . . . . . . . . 20 Dorsomedial lobe absent (Figs. 58, 59) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 Dorsomedial lobe without processes or fringes, bearing only setae and microtrichia (Fig. 55) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thienemannimyia Fittkau Dorsomedial lobe complex, including digitiform processes and/or cuticular fringes (Fig. 56) . . . 21 Abdominal tergite 8 with pair of conspicuous setal tufts; gonostylus with apical expansion forming a hood over the gonostylar tooth (Fig. 56) . . . . . . . . . . . . . . . . . . . . . . . . Helopelopia Roback Tergite 8 without tufts of setae; gonostylus without apical expansion . . . . . . . . . . . . . . . . . . . . 22 Scutal tubercle present (Fig. 22) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Conchapelopia Fittkau Scutal tubercle absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Meropelopia Roback Eye pubescent (i.e. with microtrichia between ommatidia) . . . . . . . . . . . . . . . Nilotanypus Kieffer Eye bare of microtrichia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 Midtibia with two spurs, hind tibia with one or two spurs; costa ending farther distal than CuA1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 Midtibia with one spur, hind tibia without or with one spur; costa ending level with CuA1 or more proximal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30 All tibial spurs lyrate (Fig. 40); hind tibial comb reduced, indistinct or absent . . . . . . . . . . . . . 26 At least one spur of mid- or hind tibia elongate (Fig. 39); hind tibial comb present, occasionally reduced or indistinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 Scutal tubercle present; R2 absent; costa ending clearly farther distal than CuA1 . . Larsia Fittkau Scutal tubercle absent; R2 present; costa ending little more distal than CuA1 . . . . . . . . . . . . . . . 27 Hind tibia with two spurs; gonostylus elongate, about two-thirds as long as gonocoxite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pentaneura Philippi Hind tibia with one spur; gonostylus stouter, about one-half as long as gonocoxite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Hudsonimyia Roback Spur of foretibia lyrate; mid- and hind tibiae each with one spur lyrate, the other elongate (Fig. 39); wing banded in most species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Zavrelimyia Fittkau All tibial spurs elongate; wing usually unmarked . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 Wing vein R2 present; epandrium without row of setae; parameres distinct, curving from inside gonocoxites to median and upward underneath anal point . . . . . . . . . . . . . . . Paramerina Fittkau R2 absent; epandrium with transverse row of setae; parameres indistinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Denopelopia Roback & Rutter Hind tibia with one elongate spur; epandrium with distal margin straight to weakly concave, with only few setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Monopelopia Fittkau
A group of related genera, the larvae of which are a characteristic component of streams. The group is most diverse in the Holarctic Region where at least 9 genera are recognized (Fittkau, 1962; Roback, 1971; Wiederholm, 1989). Only Thienemannimyia has been documented from Central America, but other genera are likely to have members there as well (see key couplets 21-22 and corresponding paragraphs in the Synopsis section).
448
Figs. 30.3549. Leg characters: dorsal views of apex of foretibia of (35) Pseudochironomus richardsoni (Malloch), (Nearctic, MND, fig. 29.55); (36) Chironomus sp., (Nearctic, MND, fig. 29.56); (37) Polypedilum simulans Townes, (Nearctic, MND, fig. 29.57); and (38) Nilothauma babiyi (Rempel), (Nearctic, MND, fig. 29.58); ventral views of apex of left hind tibia of (39) Zavrelimyia sp., (Nearctic, MND, fig. 29.60); (40) Larsia sp., (Nearctic, MND, fig. 29.61); (41) Nilotanypus fimbriatus (Walker), (Nearctic, MND, fig. 29.62); (42) Orthocladius consobrinus (Holmgren), (Holarctic, MND, fig. 29.64); (43) Metriocnemus sp., (Nearctic, MND, fig. 29.65); (44) Pseudochironomus richardsoni (Malloch), (Nearctic, MND, fig. 29.67); (45) Polypedilum simulans Townes, (Nearctic, MND, fig. 29.69); (46) Micropsectra sp., (Nearctic, MND, fig. 29.70); and (47) Tanytarsus sp., (Nearctic, MND, fig. 29.71); ventral views of hind fifth tarsomere and claws of (48) Phaenopsectra sp., (Nearctic, MND, fig. 29.73); and (49) Polypedilum nubeculosum (Meigen), (Holarctic, MND, fig. 29.74).
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449
31.
Hind tibia without spurs; epandrium with distal margin strongly convex, with transverse row of strong setae (Fig. 59) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Labrundinia Fittkau Wing veins r-m and R4+5 forming straight line or shallow angle (Figs. 10, 11), this angle steep only if wing reduced; calypter usually without setae (present in some Nandeva); wing cell membranes usually with setae, at least near tip of wing; TANYTARSINI . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 Crossvein r-m at steep angle to R4+5 (Fig. 9); if wing with cell membrane setae, then calypter with fringe of setae; CHIRONOMINI and PSEUDOCHIRONOMINI . . . . . . . . . . . . . . . . . . . . . . . . 49
Figs. 30.5054. Male terminalia: dorsal view of (50) Buchonomyia brundini Andersen & Sther, (Andersen & Sther, 1995, fig. 2A); (51) Djalmabatista pulchra (Johannsen), (Wiederholm, 1989, fig. 5.14E); and (52) Parochlus kiefferi (Garrett), (Holarctic, MND, fig. 29.82); ventral view of gonostylus of (53) Procladius (Psilotanypus) bellus (Loew), (Wiederholm, 1989, fig. 5.31J); dorsal view of terminalia of (54) Procladius (Holotanypus) culiciformis (Linnaeus), (Holarctic, MND, fig. 29.77). Abbreviations: dm lb, dorsomedial lobe; epand, epandrium; goncx, gonocoxite; gonst, gonostylus; gonst tth, gonostylar tooth.
450 32.
Maxillary palpus reduced, at most with three (= two free) segments; fore- and hind legs (especially tarsi) long, foretarsomere 5 without claws; antenna long, without plume, scapus and pedicel shaped similar to flagellomeres; marine. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pontomyia Edwards Palpus with more than three segments; legs and antenna not as above; various habitats . . . . . . 33 Gonocoxite appendage 1 broad basally, distal portion elongate and sickle-shaped; gonostylus with minute, toothlike apical process . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Skutzia Reiss Appendage 1 not as above; gonostylus without apical tooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
33.
Figs. 30.5559. Male terminalia (continued): dorsal views of (55) Thienemannimyia fusciceps (Edwards), (Holarctic, MND, fig. 29.80); (56) Helopelopia cornuticaudata (Walley), (Nearctic, Wiederholm, 1989, fig. 5.18H); (57) Ablabesmyia monilis (Linnaeus), (Holarctic, MND, fig. 29.78); (58) Zavrelimyia bifasciata (Coquillett), (Nearctic, MND, fig. 29.81); and (59) Labrundinia longipalpis (Goetghebuer), (Palearctic, Wiederholm, 1989, fig. 5.21G). Abbreviations: dm lb, dorsomedial lobe; gonst tth, gonostylar tooth.
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451
34. 35.
Costa ending level with CuA1 (Fig. 11) or more proximal; gonocoxite appendage 1a absent . . . 35 Costa ending distinctly distal to CuA1 (Fig. 10), or wing vestigial with venation indistinct, or gonocoxite appendage 1a broad, tonguelike . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 Pulvilli present; scutum with or without mediocentral tubercle; epandrium without paired hyaline ridges (anal crests) on or near anal point . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36 Pulvilli and scutal tubercle absent; anal crests usually present (similar to Fig. 61, 64 or 66), absent in Nandeva . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
Figs. 30.6065. Male terminalia (continued): dorsal views of (60) Rheotanytarsus calakmulensis Kyerematen & Andersen, with inset of enlarged gonocoxal appendage 2a, (Kyerematen & Andersen, 2002, figs. 35, 38); (61) Paratanytarsus sp., on right, (Nearctic, MND, fig. 29.102), and on left, anal point and part of epandrium of Paratanytarsus quadratus (Sublette), (Nearctic, Sublette, 1964, fig. 107); (62) Neozavrelia luteola (Goetghebuer), (Palearctic, Wiederholm, 1989, fig. 10.39C); (63) Caladomyia riotarumensis Reiff, (South America, Reiff, 2000, fig. 9); (64) Micropsectra sp., (Nearctic, MND, fig. 29.100); dorsal view (left) and outline of gonopod (right) with gonocoxal apodeme and paramere cross-hatched, of (65) Virgatanytarsus arduennensis (Goetghebuer), (Palearctic, Wiederholm, 1989, fig. 10.72A). Abbreviations: app, appendage; pm, paramere.
452
v app
app 1
66 Tanytarsus
67 Pseudochironomus
68 Axarus
pm app 1 app 2 goncx
gonst
pm
70 Cladopelma
71 Cryptochironomus
app 1
69 Harnischia
72 Demicryptochironomus
Figs. 30.6672. Male terminalia (continued): dorsal views of (66) Tanytarsus sp., (Nearctic, MND, fig. 29.103); (67) Pseudochironomus richardsoni Malloch, (Nearctic, MND, fig. 29.98); (68) Axarus festivus (Say), (Nearctic, MND, fig. 29.99); (69) Harnischia curtilamellata (Malloch), (cosmopolitan except Neotropical Region, MND, fig. 29.90); (70) Cladopelma spectabile (Townes), (Nearctic, MND, fig. 29.91); (71) Cryptochironomus digitatus (Malloch), (Nearctic, MND, fig. 29.85); and (72) Demicryptochironomus cuneatus (Townes), (Nearctic, MND, fig. 29.86). Abbreviations: app, appendage; goncx, gonocoxite; gonst, gonostylus; pm, paramere; v app, ventral appendage.
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453
36. 37. 38. 39. 40.
Mid- and hind tibiae with two apical spurs each (i.e. one spur on each comb) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stempellina Thienemann & Bause Mid- and hind tibiae without apical spurs . . . . . . . . . . . . . . . . . . . . . . . . Constempellina Brundin Calypter with 14 setae; epandrium without anal crests . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nandeva Wiedenbrug, Reiss & Fittkau Calypter without setae; anal crests present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38 Mid- and hind tibiae without apical spurs; anal point without spinules between anal crests; Holarctic Region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Neostempellina Reiss] Mid- and hind tibiae with one or two spurs each; anal point with spinules between anal crests . . 39 Eye hairy (i.e. with microtrichia between ommatidia). . . .Zavrelia Kieffer, Thienemann & Bause Eye bare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stempellinella Brundin Anal point tongue- or spoon shaped, usually with pair of anterodorsal crests, the latter widely separated anteriorly, tergite surface between them lacking small spines; gonostylus usually distinctly (often abruptly) narrowed in distal half; gonocoxite appendage 1a absent; appendage 2a usually with at least some lamelliform setae partially fused, the latter often forming a platelike structure distally (Fig. 60) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rheotanytarsus Thienemann & Bause Combined anal point, crest and spine configuration not as above; gonostylus usually not abruptly narrowed in distal half; gonocoxite appendage 1a present or absent; setae on appendage 2a not fused . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 Appendage 2a with lamelliform setae branched . . . . . . . . . . . . . . . . . . . Cladotanytarsus Kieffer Appendage 2a with setae unbranched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42 Antenna with 10 flagellomeres; anal point broadly rounded (Fig. 62); Holarctic, Oriental Regions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Neozavrelia Goetghebuer] Antenna with 1213 flagellomeres; anal point not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43 Mid- and hind tibiae with combs fused (Fig. 46) or narrowly separated, or rarely each comb reduced to few short spinules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44 Mid- and hind tibiae with combs clearly separate (Fig. 47) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45 Each mid- and hind tibia usually with two apical spurs, one in each comb (but spurs absent in some southern Nearctic species); epandrium with paired hyaline ridges (anal crests) at origin of anal point, these crests about as wide as long, curved, giving rounded appearance in dorsal view (Fig. 61) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paratanytarsus Thienemann & Bause Mid- and hind tibial combs usually without spurs (Fig. 46), occasionally a single spur per tibia present; anal crests, if present, longer than wide, parallel or distally converging, not giving rounded appearance (Fig. 64) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Micropsectra Kieffer, [Parapsectra Reiss; Holarctic Region] Anal point dorsal surface with two pairs of thin, posteriorly directed processes (anal point bars) (Fig. 63) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caladomyia Swedal Anal point without posteriorly directed bars (single pair of anteriorly directed bars or spines may be present) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46 Anal crests absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tanytarsus van der Wulp, in part Anal crests present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
41. 42. 43. 44.
45. 46.
454
Figs. 30.7380. Male terminalia (continued): dorsal views of (73) Microchironomus deribae (Freeman), (Palearctic, Wiederholm, 1989, fig. 10.35C); (74) Cryptotendipes casuarius (Townes), (Nearctic, MND, fig. 29.88); (75) Parachironomus hazelriggi Spies, (Nearctic, MND, fig. 29.87, as P. monochromus); (76) Paracladopelma undine (Townes), (Holarctic, MND, fig. 29.89); (77) Dicrotendipes modestus (Say), (Holarctic, MND, fig. 29.94); gonocoxal appendage 1 of (78) Einfeldia pagana (Meigen), (Holarctic, Wiederholm, 1989, fig. 10.19F); dorsal views of (79) Chironomus (Lobochironomus) montuosus Ryser, Wlker & Scholl, (Palearctic, Wiederholm, 1989, fig. 10.8D); and (80) Paratendipes albimanus (Meigen), (Holarctic, MND, fig. 29.93). Abbreviations: app, appendage; pm, paramere.
30
455
47. 48.
Anal point without clusters of small spines between anal crests . . . . . . . . . . . . . Sublettea Roback Anal point with clusters of small spines between anal crests (Figs. 65, 66) . . . . . . . . . . . . . . . . 48 Anal point dorsally with pair of anteriorly directed rods or bars that may be partially or completely fused (Fig. 65), or with short, anteriorly-directed spine in subapical position separated from spine clusters between anal crests; Holarctic, Afrotropical Regions . . . . . . . . . [Virgatanytarsus Pinder] Anal point without such processes or spines (Fig. 66). . . . . . . . . Tanytarsus van der Wulp, in part Apex of foretibia with well-developed spur with basolateral comblike teeth (Fig. 35), similar to the paired spurs on mid- and hind tibiae (Fig. 44); PSEUDOCHIRONOMINI3 . . . . . . . . . . . . . 50 Foretibia without spur or comb, at most with a spinelike extension without basolateral teeth (Figs. 3638); CHIRONOMINI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52 Wing with dark marks usually including central, irregularly shaped crossband; at least with spot around r-m and cloud below cubital fork; anal point distinctly projecting, usually with subacute tip; South America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Aedokritus Roback] Wing without dark marks; anal point absent, epandrium posteromedially at most with short, broadly truncated extension (Fig. 67) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51 Terminalia with paired or fused ventral appendages (partes ventrales) extending to posterior from anteromedial corners of gonocoxites (Fig. 67); gonocoxites without medioventral setae directed towards each other (such setae shown in Fig. 70); appendage 1a absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudochironomus Malloch Ventral appendages at most triangular in ventral view, lacking digitiform elongation to posterior; gonocoxite with irregular row of medioventral setae; appendage 1a present or absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Manoa Fittkau Antenna with 11 flagellomeres; apex of foretibia with two rounded scales, inner barely projecting beyond outer (Fig. 36) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53 Antenna with 13 flagellomeres; apex of foretibia without distinct scales, or inner scale projecting, often bearing spinelike extension (Figs. 37, 38) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79 Antepronotum medially interrupted into separate lateral lobes (Figs. 17, 18). . . . . . . . . . . . . . . 54 Antepronotum entire, although often notched medially (Fig. 16) . . . . . . . . . . . . . . . . . . . . . . . . 57 Antepronotal lobes widely separated, gap wider than deep (Fig. 18); gonocoxite appendage 1 with elongate, apically hooked distal part (similar to Fig. 81), latter bare of setae or microtrichia . . 55 Antepronotal lobes narrowly separated, gap not as wide as deep (Fig. 17); if appendage 1 with digitiform distal part, then latter carrying microtrichia and/or setae . . . . . . . . . . . . . . . . . . . . . . 56 Wing unbanded; abdominal tergites 2 or 3 to 6 with anterior, racket-shaped depressions bare of setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Glyptotendipes Kieffer Wing with medial band; abdominal tergites without depressions; Holarctic Region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .[Demeijerea Kruseman] Gonocoxite appendage 1 longer than wide (Fig. 68); epandrium without well-developed dorsomedial setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Axarus Roback Gonocoxite appendage 1 wider than long; epandrium with numerous, long dorsomedial setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xenochironomus Kieffer
49. 50.
51.
52. 53. 54. 55. 56.
Neotropical members of this tribe are in need of revision: generic limits are not all clear, and some named species may have to be transferred to other, possibly new, genera.
456 57.
Gonocoxite appendage 2 usually small, sometimes absent (Figs. 6976); if elongate then without long distal setae, if such setae present then appendages 1 and 2 both short; Harnischia complex4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58 Appendage 2 well developed, elongate cylindrical to racket-shaped, with long setae distally (Figs. 77, 7982) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72 Appendages 1 and 2 vestigial (Figs. 69, 70) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59 Appendage 1 prominent; appendage 2 variously developed (Figs. 7176) . . . . . . . . . . . . . . . . . 60 Gonostylus shorter than gonocoxite, stout, almost straight (Fig. 69) . . . . . . . .Harnischia Kieffer Gonostylus longer than gonocoxite, slender or with widened midsection, curving to median (Fig. 70) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cladopelma Kieffer Gonocoxite appendage 2 longer than anal point, subcylindrical; widespread .[Kloosia Kruseman] Appendage 2 short or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61 Appendage 2 covered by appendage 1, digitiform, without microtrichia but with 28 setae (Fig. 71); head with frontal tubercles usually present (similar to Fig. 14). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Cryptochironomus Kieffer Appendage 2, if present, not covered by superior volsella, not digitiform, and rarely with setae; frontal tubercles usually absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62 Appendage 2 absent (Figs. 7274) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63 Appendage 2 present (Figs. 75, 76) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65 Gonostylus almost straight, distally gradually narrowing (Fig. 72) . . .Demicryptochironomus Lenz Gonostylus plier-shaped, distal half curved to median and (often abruptly) narrower than basal half (Figs. 73, 74) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64 Gonostylus with apical tooth (Fig. 73). . . . . . . . . . . . . . . . . . . . . . . . . . Microchironomus Kieffer Gonostylus without apical tooth (Fig. 74) . . . . . . . . . . . . . . . . . . . . Cryptotendipes Beck & Beck Appendage 1 slender and straight or with varied apical expansion, without microtrichia but with two or three apical or subapical setae (Fig. 75); appendage 2 short, microtrichiose lobe without setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66 If appendage 1 similar to above, then extensively microtrichiose, or appendage 2 also with at least one seta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67 Setae on gonocoxite appendage 1 not arising from distinct pits; gonostylus stout except at origin, widest near middle or in distal half, its medial contour in dorsal view nearly straight; anal point slender or slightly spatulate, with or without setae, apex not curving to ventral . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Robackia Sther Appendage 1 setae usually arising from distinct pits (Fig. 75); if pits indistinct, then gonostylus partially narrow or curving, or anal point without setae or apically curved to ventral . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parachironomus Lenz Scutum with distinct central tubercle; gonocoxite appendage 1 short digitiform or tuberculate, without microtrichia; appendage 2 usually with at least one distal seta . . . . . . . . . . . . . . . . . . . 68
58. 59. 60. 61.
62. 63. 64. 65.
66.
67.
The genera treated in couplets 5871 form a diverse, worldwide array of many taxa. While the monophyly of the complex as a whole appears to be certain (Sther, 1977a, b), generic limits within it are not all clear, and continue to be challenged by additional material. New species are certain to be found in Central America, new genera are likely, and additional genera known from other regions might occur there as well.
30
457
68. 69. 70.
Scutal tubercle absent; appendage 1 oval to club- or broadly foot-shaped, at least partly covered with microtrichia (Fig. 76); appendage 2 without setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69 Epandrium with pair of digitiform, setose posterolateral projections . . . . . . . . . . Gillotia Kieffer Epandrium without posterolateral projections; widespread . . . . . . . . . . . . . . [Cyphomella Sther] Appendage 2 indicated only as weak expansion of medial gonocoxite contour; Holarctic Region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Chernovskiia Sther] Appendage 2 forming distinct lobe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70 Acrostichal setae absent; Holarctic, Afrotropical Regions . . . . . . . . . . . . . . . . [Beckidia Sther] At least few, strong acrostichals present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
Figs. 30.8182. Male terminalia (continued): dorsal views (including tergites 7 and 8) of (81) Chironomus (Chironomus) staegeri Lundbeck, (Holarctic, MND, fig. 29.96); and (82) Polypedilum walleyi Townes, (Nearctic, MND, fig. 29.97). Abbreviations: an pt, anal point; app, appendage; epand, epandrium; goncx, gonocoxite; goncx apod, gonocoxal apodeme; gonst, gonostylus; pm, paramere; tg, tergite.
458
Figs. 30.8388. Male terminalia (continued): dorsal views (left), and ventral views (right, gonocoxal apodeme and paramere cross-hatched) of (83) Thienemanniella sp., (Wiederholm, 1989, fig. 9.88E); and (84) Corynoneura sp. (Wiederholm, 1989, fig. 9.17G); dorsal view of (85) Diplocladius cultriger Kieffer, (Wiederholm, 1989, fig. 9.20E); and (86) Cricotopus laricomalis Edwards, (Holarctic, MND, fig. 29.105); dorsal view (left), and ventral view (right) of (87) Lopescladius verruculosus Sther, (Sther, 1983, fig. 2E); dorsal view of (88) Limnophyes guatemalensis Sublette & Sasa, (Sublette & Sasa, 1994, fig. 80).
30
459
71.
Epandrium with darkened bands running from anterolateral corners toward anal point, these bands in form of Y, i.e. leaving extensive lighter area between juncture of Y and anteromedial tergite margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Saetheria Jackson Epandrium without bands, I-shaped (without anterolateral arms), T-shaped (Fig. 76), or at most shallowly Y-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paracladopelma Harnisch Gonocoxite appendage 2 racket-shaped, broad distally . . . . . . . . Kiefferulus Goetghebuer, in part Appendage 2 not as broad, usually somewhat cylindrical, may be divided distally . . . . . . . . . . 73 Appendage 2 strongly bowed dorsoventrally, sometimes shallowly to deeply divided distally (Fig. 77); one or more of abdominal sternites 57 centrally often with one or more accessory setae that are flattened and blunt-tipped rather than cylindrical and acute . . . . . . . Dicrotendipes Kieffer Appendage 2 at most weakly bowed, not divided distally; sternites without accessory setae . . . 74 Appendage 1 completely digitiform, without setose basal portion . . . . Goeldichironomus Fittkau Appendage 1 with variously developed basal part that bears some long setae (Figs. 7882) . . . 75 Thoracic acrostichal setae absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76 Acrostichal setae present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77 Calypter and wing veins without setae; South America. . . . . . . . . . . . . . . . . . . . . [Pelomus Reiss] Calypter and at least some wing veins with setae; Holarctic Region. . . . . . . . [Lipiniella Shilova] Appendage 1 with digitiform process arising subapically on larger, microtrichiose base; basal part extends posterior to origin of digitiform process that is oriented obliquely to median (Fig. 78) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Einfeldia Kieffer Appendage 1 with microtrichiose base usually relatively small and confined to proximal end, digitiform process arising apically or medially; if basal part inflated, then not distinctly extending posterior to origin of digitiform process (Fig. 79) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78 Abdominal tergites without pigment pattern; wing length (from arculus, i.e. anterior branch of media, to wing tip) about 2.53 mm, area around crossvein r-m not darkened; gonocoxite appendage 1 with small, setose basal swelling and slender, acute-tipped distal process . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kiefferulus Goetghebuer, in part Usually at least some tergites with dark spot or transverse band; wing length often much greater than 3 mm, r-m area often darkened; terminalia variously shaped (e.g., Figs. 79, 81), appendage 1 often curving or hooking or with ventral apical knob . . . . . . . . . . . . . . . . . . Chironomus Meigen Apex of foretibia at most with low scale, lacking spinelike extension . . . . . . . . . . . . . . . . . . . . 80 Apex of foretibia with well-developed scale and/or spinelike extension (Figs. 37, 38) . . . . . . . 82 Calypter with fringe of setae; forefemur mediodistally with patch of posteriorly directed setae (Fig. 32) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microtendipes Kieffer Calypter without setae; forefemur without patch of posteriorly directed setae . . . . . . . . . . . . . . 81 Gonocoxite appendage 1 globose; gonostylus normally developed . . . . . . . . . .Apedilum Townes Appendage 1 digitiform; gonostylus strongly reduced . . . . . . . . . . . . . Paralauterborniella Lenz
72. 73.
74. 75. 76. 77.
78.
79. 80. 81.
460 82. 83. 84. 85. 86. 87. 88. 89. 90. 91. 92. 93.
Anal tergite with one or more dorsomedial lobes or projections bearing setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Nilothauma Kieffer5 Anal tergite without dorsomedial setose lobes or projections . . . . . . . . . . . . . . . . . . . . . . . . . . . 83 Appendage 2 fused to inner side of gonocoxite along their entire respective lengths . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fissimentum Cranston & Nolte Appendage 2 fused to gonocoxite only basally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84 Antepronotum extending clearly farther anterior than scutum (Fig. 27) . . . . . . . . Omisus Townes Antepronotum not or scarcely reaching as far to anterior as scutum (Figs. 28, 29) . . . . . . . . . . 85 Gonocoxite appendage 2a well developed (Fig. 80) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86 Appendage 2a absent or vestigial . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87 Anal point absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Beardius Reiss & Sublette Anal point present (Fig. 80) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Paratendipes Kieffer5 Anal point absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88 Anal point present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90 Gonocoxite appendage 1a present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oukuriella Epler Appendage 1a absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89 Antepronotal lobes narrowly separated medially; wing vein R1 without setae; gonostylus with blunt apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paranilothauma Soponis Antepronotum widely separated; R1 with setae; gonostylus with pointed apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neelamia Soponis Calypter bare or with at most 5 setae; abdominal segment 8 rectangular (similar to Fig. 81), not tapered anteriorly; pulvilli not bifurcate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91 Calypter usually fully fringed, if with fewer than 6 setae, then abdominal segment 8 anteriorly tapered (Fig. 82) and pulvilli with two main branches (Fig. 49) . . . . . . . . . . . . . . . . . . . . . . . . . 94 Wing marked with dark bands or spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92 Wing unmarked . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93 Abdominal tergites 27 each with mound bearing dark setae; wing with numerous dark areas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Zavreliella Kieffer Tergites without setose mounds; wing with dark bands; Nearctic Region . . . [Stelechomyia Reiss] Epandrium with pair of short, darkened, transverse bands near anterior tergite margin; appendage 1 broadly horn-shaped, without basomedial setae . . . . . . . Lauterborniella Thienemann & Bause Epandrium bands extending posterior to near anal point; appendage 1 digitiform, with basomedial group of setae; Holarctic Region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .[Pagastiella Brundin]
Watson & Heyn (1993) reported two species groups from Costa Rica that are based on Nilothauma aleta Roback and N. duena Roback, respectively. All these species lack dorsomedial anal tergite projections, and thus key to Paratendipes if the basal branch of gonocoxite appendage 1 is interpretedas Adam & Sther (1999) did for Nilothaumaas representing appendage 2a. Watson & Heyn (1993) deduce from comparisons with Robacks type material that two new genera will have to be erected.
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94. 95. 96.
Antepronotal lobes widely separated and reduced, scutum reaching far beyond their dorsal ends anteriorly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95 If antepronotal lobes reduced, then scutum not reaching beyond them . . . . . . . . . . . . . . . . . . . . 96 Combs on mid- and hind tibiae fused; gonocoxite appendage 2 slender, laterally compressed, its posterodorsal setae in linear arrangement . . . . . . . . . . . . . . . . . . . . . . . . Stenochironomus Kieffer Combs on mid- and hind tibiae separate; appendage 2 broader, subcylindrical, its setae scattered . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xestochironomus Sublette & Wirth Abdominal segment 8 tapered anteriorly (Fig. 82); gonostylus with medial submarginal setae long and more or less evenly distributed, without discrete group of subapical setae; pulvilli with two main branches (Fig. 49) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Polypedilum Kieffer Tergite 8 not tapered; medial margin of gonostylus with short setae only, often in distinct subapical group; or if setae long then not evenly distributed; pulvilli simple (Fig. 48) . . . . . . . . . . . . . 97
Figs. 30.8993. Male terminalia (continued): dorsal view (left), and ventral view (right; gonocoxal apodeme and paramere stippled) of (89) Irisobrillia longicosta Oliver, (Oliver, 1985, fig. 32); (90) Parakiefferiella sp., (Nearctic, MND, fig. 29.110); (91) Orthocladius decoratus (Holmgren), (Holarctic, MND, fig. 29.107); (92) Antillocladius pluspilalus Sther (Wiederholm, 1989, fig. 9.4E); dorsal view (left) and ventral view (right, gonocoxal apodeme and paramere cross-hatched) of (93) Parametriocnemus lundbeckii (Johannsen), (Wiederholm, 1989, fig. 9.56F).
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Figs. 30.9498. Male terminalia (concluded): dorsal views (left) and ventral views (right, gonocoxal apodeme and paramere cross-hatched) of (94) Mesosmittia tora Sther, (Nearctic, Wiederholm, 1989, fig. 9.46D); and (95) Bryophaenocladius flavoscutellatus (Malloch), (Nearctic, Wiederholm, 1989, fig. 9.9F); dorsal views of (96) Smittia sp., (Holarctic, MND, fig. 29.109); and (97) Diplosmittia carinata Edwards, (Nearctic, Wiederholm, 1989, fig. 9.21D); dorsal view (left) and ventral view (right, gonocoxal apodeme and paramere cross-hatched) of (98) Pseudosmittia forcipata Goetghebuer, (Wiederholm, 1989, fig. 9.71E).
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97. 98. 99.
Wing with setae in cell membranes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Phaenopsectra Kieffer Wing cell membranes bare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98 Scutal tubercle present (Fig. 28); gonostylus distally flattened, movable against gonocoxite; tibiae often banded; widespread. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Stictochironomus Kieffer] Scutal tubercle absent; gonostylus rigidly fused to gonocoxite; tibiae not banded . . . . . . . . . . . 99 Gonocoxite appendage 1 with strong seta in lateral subapical position. . . Endotribelos Grodhaus Appendage 1 with setae only on anteromedial basal part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100
100. Foretarsus without beard of elongate setae; mid- and/or hind legs usually with fewer than 2 spurs per tibia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Tribelos Townes Foretarsus with beard; if beard setae relatively short (some Endochironomus), then mid- and hind tibiae usually with two spurs each . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
101. Foretibia with one apical scale ending in spinelike extension; palpus not reduced (similar to Fig. 14) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Endochironomus Kieffer Foretibia without spine; segments 35 shortened; Nearctic Region . . . . . . . [Hyporhygma Reiss]
102. Wing with veins R1 and R2+3 short and thick, often fused with C in thick clavus ending before midlength of wing; R4+5 weak (Fig. 6) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103 Wing with veins R1, R2+3 (if present) and R4+5 narrow, elongate, separate until their respective meeting with C beyond midlength of wing (Figs. 5, 7, 8) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
103. Genital segment internally with gonocoxal apodeme dilated, inverted V- or U-shaped, with no trace of oral projections (Fig. 84); eye bare of microtrichia between ommatidia; foretrochanter with dorsal keel (Fig. 34); apex of hind tibia enlarged (Fig. 33) . . . . . . . . Corynoneura Winnertz Gonocoxal apodeme narrow, straight or convex; anterolateral corners slightly to strongly projecting to oral (Fig. 83); eye bare or with microtrichia between ommatidia; dorsal keel of foretrochanter at most moderately developed; apex of hind tibia at most slightly enlarged . . . . . . . . 104
104. Eye with microtrichia between ommatidia; foretrochanter without dorsal keel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thienemanniella Kieffer Eye bare; foretrochanter with moderately developed dorsal keel . . . . . . . . . . . . . . . . . . . . . . . 105
105. Antenna with 13 flagellomeres; abdominal tergites 25 with distinct central cluster of setae on circular mound . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Tempisquitoneura Epler Antenna with 1112 flagellomeres; tergites with 48 setae in transverse row, cuticula without central mound . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Onconeura Andersen & Sther
106. Hind tibia without apical comb of stiff setae; palpus with at most three (= two free) distinctly separate segments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107 Hind tibial comb present (Figs. 42, 43), or palpus with four or five (three or four free) segments (Figs. 12, 13), or both these conditions met . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108
107. Antenna with at most 9 flagellomeres, sparsely setose; wing broad (Fig. 5); tarsomeres 5 weakly cordiform or bilobed; immature stages in marine habitats. . . . . . . . . . . . . . . . . . . .Clunio Haliday Antenna with 13 flagellomeres, fully plumed; wing slender; all tarsomeres cylindrical; immatures in freshwater . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Symbiocladius Kieffer
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108. Eye hairy (i.e. with microtrichia longer than height of ommatidia). . . . . . . . . . . . . . . . . . . . . . 109 Eye bare or pubescent (microtrichia, if present, at most as long as ommatidia height). . . . . . . 118
109. Gonostylus bifurcate at base (Fig. 85) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 110 Gonostylus simple, at most with lobe or projection not branching off at base of gonostylus . . . 111
110. Antenna with terminal flagellomere longer than remaining flagellomeres combined (antennal ratio AR greater than 1.0); wing with cubital fork level with crossvein r-m . . . . . . Diplocladius Kieffer Terminal flagellomere much shorter than combined length of remaining flagellomeres (AR less than 0.5); cubital fork farther distal than r-m; South America, Australia . . . . . . .[Stictocladius Edwards]
111. Thoracic dorsolateral setae short and/or decumbent, often arising from pale areas on scutum . . . 112 Dorsocentral setae elongate and erect, scutum usually without pale areas around origins of setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113
Figs. 30.99101. Pupae: lateral views of entire pupa and dorsal views of abdominal segments 79 of (99) Procladius sp., (Nearctic, MND, fig. 29.138); (100) Chironomus sp., (Nearctic, MND, fig. 29.139); and (101) Orthocladius sp., (Nearctic, MND, fig. 29.140).
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112. Wing membrane without dark spots or bands; terminalia as in Fig. 86 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Cricotopus van der Wulp, in part Wing membrane with dark spots or bands . . . . . . . . . . . . . . . . . Cricotopus van der Wulp, in part; Oliveiriella Wiedenbrug & Fittkau
113. All tarsomeres 4 cordiform (similar to Fig. 31); microtrichia of eye as long as ommatidia height; gonocoxite with or without long extension past origin of gonostylus (Fig. 87) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lopescladius Oliveira Tarsomeres 4 cylindrical; microtrichia of eye longer than ommatidia height; gonocoxite not extended past origin of gonostylus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 114
114. Anal point absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115 Anal point present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116
115. R4+5 ending at level of CuA1 or more proximal; thoracic acrostichal setae absent; eye without dorsomedial extension . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eukiefferiella Thienemann, in part R4+5 ending more distal than CuA1; acrostichals present; eye with short dorsomedial extension . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paratrichocladius Santos Abreu
116. Antenna with strong subapical seta (Fig. 15) . . . . . . . . . . . . . . . . . . . . . Smittia Holmgren, in part Antenna without strong subapical seta. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
Figs. 30.102104. Fourth instar larvae: lateral views of (102) Ablabesmyia sp., (Nearctic, MND, fig. 29.114); (103) Chironomus sp., (Nearctic, MND, fig. 29.115); and (104) Pseudosmittia sp., (Nearctic, MND, fig. 29.116). Abbreviation: prcerc, procercus.
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117. Thorax with at most six acrostichal setae concentrated at midscutum (Fig. 24); eye protruding (Fig. 12); abdominal tergites medially with setae in one or more rows across midtergite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nanocladius Kieffer Acrostichals usually more numerous and starting near antepronotum; eye not protruding (similar to Fig. 13); abdominal tergites medially with setae in group, and occasionally with anterior and/ or posterior transverse row . . . . . . . . . . . . . . . . . . . . . . . Rheocricotopus Thienemann & Harnisch
118. Thorax with setae on posterior anepisternum and epimeron, usually also on katepisternum and dorsal parts of antepronotum; often some humeral and/or prescutellar setae lanceolate (Fig. 21); terminalia as in Fig. 88 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Limnophyes Eaton Posterior anepisternum, epimeron and dorsal parts of antepronotum bare; no lanceolate setae on scutum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119
119. Thoracic acrostichal setae absent, but tuft of microtrichia may be present at midscutum. . . . . 120 Acrostichals present, but may be few, short and concentrated at midscutum . . . . . . . . . . . . . . 124
120. Membrane of some wing cells with setae; terminalia as in Fig. 89 . . . . . . . . . . .Irisobrillia Oliver Wing with setae only on veins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121
Figs. 30.105106. Fourth instar larvae (concluded): ventral views of head capsule of (105) Ablabesmyia sp., (Nearctic, MND, fig. 29.117); and (106) Chironomus sp., (Nearctic, MND, fig. 29.119). Abbreviations: ant, antenna; epiphar b, epipharyngeal bar; eyesp, eyespot; hyphar pct, hypopharyngeal pecten; hyps, hypostoma; lig, ligula; md, mandible; pagl, paraglossa; palb plt, paralabial plate; premd, premandible; tm, torma.
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121. Tarsomeres 4 more or less cordiform (Fig. 31); anal point absent. . . . . . . . Cardiocladius Kieffer Tarsomeres 4 cylindrical; anal point generally present (Figs. 90, 91) . . . . . . . . . . . . . . . . . . . . 122
122. Calypter without setae; scutum with medial tuft of microtrichia (Fig. 25); terminalia as in Fig. 90 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parakiefferiella Thienemann Calypter with fringe of setae; scutum without medial microtrichial tuft (Fig. 23) . . . . . . . . . . 123
123. R4+5 ending farther distal than CuA1; endophallus with sclerotized spines (virga) present (shown in Fig. 91 underneath midpoint of gonocoxal apodeme) or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Orthocladius van der Wulp, in part R4+5 ending level with CuA1 or more proximal; virga absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Synorthocladius Thienemann, in part6
124. Thoracic acrostichal setae all scalpellate (Fig. 26), or some scalpellate acrostichals present near midscutum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125 Acrostichals straight or hooked, but all hairlike . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 128
125. Calypter bare of setae; costa with long extension beyond R4+5; acrostichal setae present only at some distance from antepronotum; endophallus without sclerotized spines (virga); gonostylar tooth simple or distally dentate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Compterosmittia Sther If calypter bare, then some acrostichals positioned near antepronotum; costal extension short or long; virga usually present (as in Fig. 92, and Fig. 91 underneath midpoint of gonocoxal apodeme); gonostylar tooth simple, not dentate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 126
126. Two small, scalpellate acrostichal setae present at midscutum; R4+5 ending level with CuA1 or more proximal; virga absent. . . . . . . . . . . . . . . . . . . . . . . . . Synorthocladius Thienemann, in part Acrostichals more numerous, all scalpellate or some anterior ones hair-like, their rows beginning near antepronotum or near mid-scutum; R4+5 ending farther distal than CuA1; virga present or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127
127. Wing with coarse punctation (microtrichia visible at 60x magnification); cell membranes without setae; anal point nearly parallel sided, apically rounded, hyaline, placed at end of medial, setose ridge of epandrium; virga absent; Holarctic Region, Australia . . . . . . .[Paralimnophyes Brundin] Wing with fine punctation; cell membranes near wing apex often with setae; anal point long, pointed with strong lateral setae; virga present (Fig. 92) or absent . . . . . . . .Antillocladius Sther
128. Membrane of some wing cells with setae, at least near wing tip . . . . . . . . . . . . . . . . . . . . . . . . 129 Wing with setae only on veins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131
129. R4+5 always and C usually ending more proximal than CuA1 . . . Paraphaenocladius Thienemann R4+5 usually and C always ending level with CuA1 or farther distal . . . . . . . . . . . . . . . . . . . . . 130
130. Wing vein CuA2 straight; head and thorax with numerous strong setae: inner verticals, supraalars (= prealars, not supraalars of Sther 1980) and scutellars bi- to multiserial . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Metriocnemus van der Wulp CuA2 distinctly curved; inner vertical, supraalar and scutellar setae usually uniserial; terminalia as in Fig. 93 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parametriocnemus Goetghebuer
Keyed here because the two small acrostichal setae at mid-scutum rub off easily; see also couplet 126.
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131. Calypter with at least 1 seta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132 Calypter bare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
132. Epandrium with medial longitudinal, strongly elevated, sclerotized ridge, without true anal point (Fig. 94) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mesosmittia Brundin If epandrium with elevated ridge, then anal point present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133
133. Anal point absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134 Anal point present (Figs. 91, 95) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136
134. Acrostichals weak, often scarcely visible; dorsocentrals uniserial; R4+5 ending level with CuA1 or more proximal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eukiefferiella Thienemann, in part Acrostichals strong; dorsocentrals partially bi- to triserial; R4+5 ending more distal than CuA1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
135. Scutellar setae uniserial; mid- and hind tarsomeres 1 with sensilla chaetica (short setalike structures with tips bent at right angle) in row along distal one-fifth of tarsomere . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Orthocladius van der Wulp, in part Scutellars biserial; mid- and hind tarsomeres 1 without sensilla chaetica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lipurometriocnemus Sther
136. Anal point hyaline, semicircular to subtriangular, projecting from setose area on posterior margin of epandrium (Fig. 95); wing membrane with coarse punctation (microtrichia visible at 60x magnification); thoracic acrostichal setae decumbent but strong . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Bryophaenocladius Thienemann, in part Anal point usually strong and with lateral setae, triangular and pointed (Fig. 91), or parallel sided with rounded apex; wing membrane with fine to moderate punctation; acrostichals often weak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Orthocladius van der Wulp, in part
137. Eye pubescent with microtrichia; antenna with strong subapical seta (Fig. 15); acrostichals absent; anal point slender, elongate (Fig. 96) . . . . . . . . . . . . . . . . . . . . . . . . . . . Smittia Holmgren, in part Eye bare; acrostichals present; anal point, if present, usually broadly triangular or short (Figs. 95, 97, 98) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138
138. Acrostichal setae decumbent but strong, their rows beginning close to antepronotum; terminalia as in Fig. 95 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Bryophaenocladius Thienemann, in part Acrostichals weak, usually few and limited to midscutum . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139
139. Gonostylus deeply split into two long branches; anal point distinctly projecting from posterior margin of epandrium, triangular with broad base and rounded tip, with microtrichia and lateral setae (Fig. 97) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diplosmittia Sther Gonostylus simple, or with short to moderately elongate heel; anal point, if present, originating in more dorsal position, rarely extending beyond posterior margin of epandrium, variously shaped but generally smaller, with microtrichia but without strong setae (Fig. 98) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudosmittia Edwards
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469 Bryophaenocladius Thienemann. Only one named species of this widespread and species-rich genus has been recorded from Central America: B. carus (Roback), described from Panama. However, undescribed species have been found in southern Mexico (Contreras Ramos & Andersen, 1999), and many more are expected to occur in the region. Adult males of North American Bryophaenocladius are keyed in Wang et al. (2004). Larvae of most species live in semi-terrestrial or terrestrial habitats. Buchonomyia Fittkau. There is a single record of Buchonomyia from Central America: B. brundini Andersen & Sther from Costa Rica (Andersen & Sther, 1995). The immature stages of this species, its habitat and ecology are unknown. Caladomyia Swedal. One named species is recorded: C. pistra Sublette & Sasa from Guatemala. However, based on what is known from adjacent areas to the south and north (Swedal, 1981; Reiff, 2000; Trivinho-Strixino & Strixino, 2000, 2003), the genus is expected to be widely distributed and species rich in Central America. Larvae are found on submerged aquatic macrophytes. Cardiocladius Kieffer. This genus has been recorded from Costa Rica (Watson & Heyn, 1993) and Mexico (Vockeroth, 1949). Larvae live in fast-flowing water, often associated with Simuliidae. Predation on the latter has been suggested but is unproven. Chironomus Meigen. This is one of the most widely distributed, common and species-rich chironomid genera. Its members are grouped in several subgenera differing in adult male genitalia structure. Only five species in C. (Chironomus) are well documented from Central America (Fittkau, 1965; Wlker et al., 1989; Sublette & Sasa, 1994; Spies & Reiss, 1996)but see genus Einfeldia belowseveral others are named but unrecognizable from the original descriptions (Spies & Reiss, 1996). Many more are expected to be found, the low present count being due to the amount of effort necessary to identify the species (e.g., Spies, 2000b; Epler, 2001; Spies et al., 2002). Larvae inhabit soft sediments and other deposits of organic matter in all kinds and sizes of water bodies. Some species require relatively pristine freshwater and others are tolerant of acidic or adapted to brackish water. Many thrive in nutrient-enriched, low-oxygen or even polluted habitats. High survival success under conditions restricting the numbers of other organisms often leads to enormous populations whose emerging and swarming adults can interfere with human affairs. Cladopelma Kieffer. The one species recorded, Cladopelma forcipis (Rempel), occurs throughout Central America and adjacent mainland regions (see Sublette & Sasa, 1994). Larvae of Cladopelma are found in sandy or muddy sediments of lakes and rivers.
Synopsis of the fauna

This section offers remarks not only on genera for which inclusion is based on published reports or collection material, but also on a few that have not been recorded in Central America but are considered highly likely to occur there because they are known from adjacent regions both to the north and south. For additional information on genera mentioned in the Synopsis or Key see Wiederholm (1989), Spies & Reiss (1996), or Epler (2001). Some specimens noted below are in the Zoologische Staatssammlung Mnchen (ZSM) or the Zoological Museum, University of Bergen (ZMUB). Ablabesmyia Johannsen. This widespread and speciesrich genus is divided into four subgenera (e.g., Wiederholm, 1989), all of which have representatives in the Neotropical Region. Two named species have been recorded from Central America: A. cinctipes (Johannsen) from Mexico, Belize and Guatemala; and A. costarricensis (Picado), described as larvae, pupae and adult females obtained from bromeliad phytotelmata. Watson & Heyn (1993) list adults of three unnamed species from Costa Rica. Ablabesmyia larvae live in a wide variety of habitats. Alotanypus Roback. One of the two named American species, A. venustus (Coquillett), is distributed from western North America at least to Costa Rica (Roback, 1971; Watson & Heyn, 1993; Sublette & Sasa, 1994). The larval ecology has not been reported, but a broad range of conditions appear to be tolerated (e.g., Roback, 1971: adults from diverse waters; Sublette & Sasa, 1994: coffee pond; Spies, 2000b: urban stream). Antillocladius Sther. Six named species have been described or recorded from southern Mexico, Nicaragua or Costa Rica (Mendes et al., 2004; this includes keys). The known larvae are semiaquatic, living in spring seeps, the margins of streams, etc. Apedilum Townes. Two named species have been recorded: A. elachistum Townes (widespread), and A. subcinctum Townes (Guatemala, southern Mexico). There are several undescribed Neotropical species (see Epler, 1988a). Apedilum larvae are found on aquatic macrophytes in fresh or brackish water. Under favorable conditions, emergence may take place all year, with life cycles as short as one week (Nolte, 1995). Axarus Roback. One named species is recorded: A. rogersi (Beck & Beck) from Costa Rica, Mexico, and Nicaragua. More species are likely to occur (e.g., see Andersen & Mendes, 2002a). Larvae are found in soft sediments and peaty detritus. Beardius Reiss & Sublette. This is a widespread genus, with five named species recorded (see Reiss & Sublette, 1985; Andersen & Sther, 1996; Jacobsen & Perry, 2000). Larvae are found in lentic and lotic habitats.
470 Cladotanytarsus Kieffer. This genus is common and species rich in the Nearctic Region, but there are only a few records of unnamed species from south of the USA (Spies & Reiss, 1996). Larvae may be found in a variety of lentic and lotic habitats, including brackish water and hot springs. Clinotanypus Kieffer. This genus has been reported only as unnamed adults from Mexico (Contreras Ramos & Andersen, 1999), but several species are known from the southern USA and Brazil, respectively. Two subgenera are recognized, but C. (Aponteus) Roback is known from the female only. See Roback (1971) and Oliveira (1953). The larvae inhabit soft sediments of lakes and pool areas of streams. Clunio Haliday. Although this genus has not yet been recorded from Central America, it is present in all adjacent regions and worldwide. These flies are marine, usually found in intertidal habitats, seawater inlets, etc., but Schrer & Epler (2007) have found larvae among algae on the backs of sea turtles caught 40 miles off a Caribbean coast. See Stone and Wirth (1947). Adult females are vermiform, wingless, with strongly reduced legs. Males skate over water and substrate surfaces, and mate with females often while the latter are still teneral or even pharate. In many species, emergence is highly synchronized with the lunar cycle, and adults survive for little longer than the period between low tides. Coelotanypus Kieffer. Eight named species of this widespread and species-rich genus are known from Central America (Spies & Reiss, 1996; Andersen et al., 2000), but much Neotropical material is awaiting description. See Roback (1965, 1971). The larvae inhabit soft sediments in lakes and slow-flowing running waters. Compterosmittia Sther. Two species are known from Central America: C. aberrans Mendes, Andersen & Sther from Costa Rica (Mendes et al., 2004; this includes keys), and C. cf. nerius (Curran) from Nicaragua (ZSM collection). The only known larva in the genus (of an Oriental species) was taken from water held by a pitcher plant. Conchapelopia Fittkau. This widespread genus has not been recorded from Central America, but larvae with a 3-segmented b-sensillum on the maxillary palpus, thus possibly belonging to Conchapelopia or Helopelopia, have been found in Costa Rica (C.N. Watson, unpublished). See Roback (1971), Bilyj (1985). Larvae live in lotic and lentic waters, usually preferring oxygen-rich habitats. Constempellina Brundin. Although not known from Central America, this genus has been recorded in North and South America. Larvae live in streams and build transportable sand cases. Corynoneura Winnertz. Two named species of this widespread and species-rich genus, C. ferelobata Sublette & Sasa and C. hirvenojai Sublette & Sasa, have been described from Guatemala (Sublette & Sasa, 1994). In addition, the genus
has been recorded from Costa Rica (Coffman et al., 1993) and southern Mexico (Contreras Ramos & Andersen, 1999). Larvae occur in all kinds of permanent aquatic habitats. Cricotopus van der Wulp. This is one of the most widely distributed, common and species-rich chironomid genera. Six named species have been recorded in Central America (Sublette & Sasa, 1994; Spies & Reiss, 1996; Contreras Ramos & Andersen, 1999), and many more are expected to occur there. Unnamed or new material has been recorded from Costa Rica (Watson & Heyn, 1993), Nicaragua (Palomki, 1987; ZSM collection), and Panama (ZSM collection). Adults with banded wings are generically inseparable from those of Oliveiriella. Larvae are found in all types of freshwater, and some species tolerate elevated salinity levels. Associations with plants are common, for instance with benthic or floating algae, or as miners in macrophytes (e.g., causing damage in rice fields). Cryptochironomus Kieffer. In this genus, a number of species can be difficult to separate in the adult stage; pupae are often easier to identify (Mason, 1986). Only a few records exist from Central America (Spies & Reiss, 1996). Larvae occur in lakes, rivers and streams and are often associated with sandy sediments. Cryptotendipes Beck & Beck. There is a single record of unnamed larvae of this genus from Nicaragua (Palomki, 1987). Larvae are found in sandy or muddy sediments of lakes and rivers. Demicryptochironomus Lenz. Although not known from Central America, this genus has been recorded in North and South America. Larvae are found in soft or sandy sediments of lakes and rivers. Denopelopia Roback & Rutter. The single described species, D. atria Roback & Rutter, 1988, is known from Florida and Costa Rica (Epler, 2001). Unnamed material at ZSM includes an adult male from Panama (Spies & Reiss, 1996). Larvae have been found in ditches and ponds with dense macrophyte stands, and can tolerate low dissolvedoxygen and high iron concentrations. Dicrotendipes Kieffer. Six named species of this widespread and common genus are known from Central America. See Epler (1988b, which includes keys) and Epler (1996b). Larvae are found on plants and sediments in lentic and lotic waters. Diplocladius Kieffer. The only species described in this genus, D. cultriger Kieffer, has been recorded from Nicaragua (Bylmakers and Sobalvarro, 1988). The larvae inhabit springs and other small bodies of water. Diplosmittia Sther. Two named species of Diplosmittia are recorded, D. beluina Andersen and D. forficata Andersen, both described from Costa Rica (Andersen, 1996, which in-
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471 Mexico (Andersen et al., 2000). See also Serrano & Nolte (1996). The larvae of most species live in soft sediments of springs and montane lakes. Gillotia Kieffer. This genus has not been recorded from Central America, but is known from the southern USA and Brazil. Its few member species are rare, and no information is available on their ecology. Glyptotendipes Kieffer. No described species are recorded from Central America, but unnamed ones have been collected from Costa Rica (Watson & Heyn, 1993) and southern Mexico (Contreras Ramos & Andersen, 1999). Larvae are found in lakes, ponds, rivers and streams, often in eutrophic conditions, and some are miners in aquatic plants or immersed wood. Goeldichironomus Fittkau. This genus is limited to the New World, where its members are commonly encountered in subtropical and tropical areas. All four named species known from Central America are widely distributed there and beyond. One undescribed species is known from Costa Rica (J.H. Epler collection), and more species are expected to occur in the area. See Reiss (1974), Trivinho-Strixino & Strixino (1998), and references in these. Two species, G. amazonicus (Fittkau) and G. holoprasinus (Goeldi), are successful in waters impacted or created by man, and apparently through human transport have reached the southern USA and Hawaii, respectively. Goeldichironomus larvae occur on sediments and plants, often in eutrophic conditions. Harnischia Kieffer. In Central America, this genus has been recorded only as pupae from Costa Rica (Watson & Heyn, 1993). Larvae are found in sediments of lakes and rivers. Helopelopia Roback. This Nearctic genus has not been recorded from Central America, but larvae with a 3-segmented b-sensillum on the maxillary palp, thus possibly belonging to Conchapelopia or Helopelopia, have been found in Costa Rica (C.N. Watson, unpublished). See Roback (1971). The known larvae live in streams. Hudsonimyia Roback. Sublette & Sasas (1994) relegation of Hudsonimyia to subgenus status in Pentaneura Philippi is not followed here. The single species known from Central America, H. vittaria (Sublette & Sasa), comb. nov., was described from Guatemala. See also the references cited in Sublette & Sasa (1994). Larvae were found in special hygropetric habitats: shallow water flowing slowly over granite outcrops covered with algae, moss and detritus. Irisobrillia Oliver. The single included species, I. longicosta Oliver, 1985, has been recorded in Central America from Costa Rica and Nicaragua (Watson & Heyn, 1993; Spies & Reiss, 1996). The immature stages and habitat are unknown.
cludes a key). The genus has been recorded in southern Mexico (Contreras Ramos & Andersen, 1999) and Nicaragua (ZSM collection). The immature stages and habitat are unknown. Djalmabatista Fittkau. Although this genus is diverse in the Neotropical Region, only one named species has been recorded from Central America, the widespread D. pulchra (Johannsen). Two undescribed species are known from Costa Rica (Watson & Heyn, 1993), and more are likely awaiting discovery. Descriptions are listed in Spies & Reiss (1996), Oliveira & Carraro (1997), and Roback (1980). Larvae occur in lentic and lotic depositional habitats, and appear to prefer low-alkalinity, weakly acidic water. Einfeldia Kieffer. This genus requires revision. Some species described earlier in Einfeldia have been transferred to Chironomus, but generic distinction and placement can be difficult. According to the criterion proposed in Wiederholm (1989; see also the Key above), the single named species known from Central America, E. atitlanensis Sublette & Sasa, 1994, belongs in Chironomus (Lobochironomus) Ryser, Wlker, & Scholl. On the other hand, Townsia subaberrans (Walley), so far known only from Guyana, but possibly more widely distributed, probably belongs in Einfeldia. Maes (1990) gives a generic record from Nicaragua. Larvae of Einfeldia are usually found in sediments in eutrophic conditions. Endochironomus Kieffer. From Central America, there is a single, tentative record of E. subtendens (Townes) from southern Mexico (Spies & Reiss, 1996). See Grodhaus (1987), which includes keys. Larvae live in lentic and lotic waters, on sediments and plants, or as miners in macrophyte leaves. Endotribelos Grodhaus. Two named species are known from Guatemala or Costa Rica (Spies & Reiss, 1996), and Watson & Heyn (1993) reported three undetermined species from Costa Rica. See Grodhaus (1987), which includes keys. The larvae of some species are miners in aquatic plants. Eukiefferiella Thienemann. This genus is widespread and species-rich in other parts of the world, but rarely recorded from the Neotropical Region. Watson & Heyn (1993) report larvae from Costa Rica. Habitats are primarily lotic. Fissimentum Cranston & Nolte. This is a tentative first record for Central America, based on larvae from Costa Rica (J.H. Epler collection). Although a number of similar larvae fit the generic diagnosis of Cranston & Nolte (1996), other life stages are known for only one Fissimentum species. Cranston & Spies (2000) and Epler (2001) suggest the possibility that not all species with such larvae are congeneric. Specimens were taken from fine sediments in a variety of water bodies. Fittkauimyia Karunakaran. Found worldwide in tropical and subtropical regions, this genus is recorded in Central America from Costa Rica (Watson & Heyn, 1993) and
472 Kiefferulus Goetghebuer. There is a single record of unnamed material of this genus from southern Mexico (Andersen et al., 2000). Larvae are found in sediments in eutrophic or oligotrophic habitats. Labrundinia Fittkau. Most species of this worldwide genus are found in the Neotropical to southern Nearctic Regions; many are known as larvae only (Roback, 1987; Epler, 2001). The single record from Central America (Watson & Heyn, 1993: larvae from Costa Rica) poorly reflects the diversity expected there. See Roback (1971, 1987). Larvae live in small bodies of standing water and in running waters. Larsia Fittkau. Three named species of this worldwide genus have been found in Central America (Sublette & Sasa, 1994, which includes a key). Larvae live in various habitats, including boggy pools, wet Sphagnum stands, and hot springs. Laurotanypus Oliveira, Messias & da Silva-Vasconcelos. The only species so far included in this genus, L. travassosi Oliveira, Messias & da Silva-Vasconcelos, 1992 from Brazil, resembles some species currently placed in Procladius (Psilotanypus) Kieffer. Type material should be compared with P. (Ps.) stroudi Roback, 1982a, to investigate possible synonymy. Procladius (Ps.) etatus Roback, 1982a, known from Costa Rica (Watson & Heyn, 1993), possesses the adult scutal tubercle proposed as diagnostic for Laurotanypus, but its wing is darkened only around crossvein r-m. Generic placement of these species remains tentative until their immature stages are discovered. Lauterborniella Thienemann & Bause. This record is based on adult males from Panama (Spies & Reiss, 1996) of a single species, L. agrayloides (Kieffer). Larvae build silken transportable cases and live amongst vegetation in ponds and lakes. Limnophyes Eaton. Two named species of this widespread and species-rich genus have been described from Guatemala (Sublette & Sasa, 1994). See also Sther (1990a, b). Larval habitats in the genus range from aquatic to terrestrial. Lipurometriocnemus Sther. The only named species included, L. glabalus Sther, was described from St. Vincent (Sther, 1981). Cranston & Oliver (1988) report probably conspecific adult males from Costa Rica. The immature stages and habitat are unknown. Lopescladius Oliveira. One species of this New World genus, L. (Lopescladius) verruculosus Sther, has been described from southern Mexico (Sther, 1983). The current subgenus L. (Cordiella) Coffman & Roback, in which the gonocoxite is not posteriorly extended, and which may also occur in Central America, probably merits separate genus rank (P.S. Cranston, personal communication). Pupae of Lopescladius have been recorded from Costa Rica (Watson & Heyn, 1993). Larvae live in sandy sediments of flowing waters.
Manoa Fittkau. Although this genus is not yet known from Central America, it occurs in southern Florida and Brazil. See Fittkau (1963), and Jacobsen & Perry (2002). Larvae live in shallow-water habitats occurring seasonally next to larger bodies of water with considerable fluctuations in water level. Meropelopia Roback. Sublette & Sasa (1994) described a pupa from Guatemala that could belong to this Holarctic genus. An identical pupa has been found in Costa Rica (C.N. Watson, unpublished). Mesosmittia Brundin. Five named species have been described from Central America (see Andersen & Mendes, 2002b). The specific nature of the larval habitat is unknown (Wiederholm, 1989; Epler, 2001). Metriocnemus van der Wulp. Four named species of this widespread and species-rich genus have been described from Central America (Picado, 1913; Sublette & Sasa, 1994). A key for the Holarctic Region is given in Sther (1995). The larvae live in a variety of mostly small, sometimes semi-terrestrial habitats, in Central America probably often in phytotelmata (e.g., see Picado, 1913). Microchironomus Kieffer. A single named species, M. nigrovittatus Malloch, has been recorded from southern Mexico (Andersen et al., 2000). Larvae live in a variety of habitats, including brackish water. Micropsectra Kieffer. This genus is common and speciesrich in North America, but rarely recorded in the Neotropical Region, where only one named species is known, M. atitlanensis Sublette & Sasa, 1994 (from Guatemala). Watson & Heyn (1993) report unnamed adults from Costa Rica. Larvae are found in a variety of situations ranging from hygropetric habitats or temporary pools to soft sediments in small rivers and lakes. The genus Parapsectra Reiss cannot be reliably separated from Micropsectra in the adult stage. Pupae are necessary for generic identification. Microtendipes Kieffer. No named species are identified from Central America, but larvae have been collected in Costa Rica (Watson & Heyn, 1993) and Nicaragua (Palomki, 1987). The common habitat is sediments in larger lentic and lotic waters. Monopelopia Fittkau. In the New World, this genus is known from North America, South America, and the Caribbean (Roback, 1986a; Serpa-Filho & Oliveira, 1997; Epler & Janetzky, 1999; Mendes et al., 2003). It is thus expected to occur in Central America as well. The larvae live in small bodies of open water; some species apparently specialize in phytotelmata. Naelotanypus Roback. The only included species, N. viridis Roback, is known from Colombia (Roback, 1982b) and Surinam. Other life stages have not been associated di-
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473 Onconeura Andersen & Sther. One named species has been recorded, O. semifimbriata (Sther, 1981) from Costa Rica, Guatemala and Mexico (Andersen & Sther, 2005). At the type locality on St. Vincent, the larvae were found in the benthos of a river. Orthocladius van der Wulp. This genus is widespread, with several morphologically diverse subgenera and many species at least in the Holarctic Region, but rarely reported from the New World tropics. Watson & Heyn (1993) list adults of the subgenus O. (Eudactylocladius) Thienemann from Costa Rica. Keys are presented in Soponis (1977, 1990) and Cranston (1999). Larvae are found in all kinds of habitats, including hygropetric, temporary, or semiterrestrial ones. Oukuriella Epler. This genus is limited to the Neotropical Region, but widespread there, with four named species known from Costa Rica or southern Mexico. See Epler (1986a, 1996a) and the various papers by Messias and coauthors. Larvae have been found on immersed wood (Messias et al., 2000a) and in freshwater sponges (Roque et al., 2004). Parachironomus Lenz. This genus is widespread and species rich, with five named species known from Central America, but many more from adjacent regions. Keys to species are given in Spies et al. (1994) and Spies (2000a). Larvae are found in a variety of lentic and lotic habitats. Their feeding ecology varies among species, from opportunistic gathering to mining in Bryozoa or macrophytes, or even to ectoparasitism on other invertebrates. Paracladopelma Harnisch. A single record exists of adults of a tentatively identified species from Costa Rica (Watson & Heyn, 1993). See Jackson (1977), and also under Saetheria below. Larvae are found in sandy sediments of lakes, ponds, rivers and streams. Parakiefferiella Thienemann. Although not recorded from Central America, this genus is known from adjacent areas to the north and south. See Wiedenbrug & Andersen (2002). Larvae are found in lotic and lentic habitats. Paralauterborniella Lenz. The only currently included species, P. nigrohalteralis (Malloch), has been recorded from Costa Rica and Nicaragua (see Spies & Reiss, 1996). Its larvae occur in streams and rivers. Paramerina Fittkau. Two named species have been recorded: P. fasciata Sublette & Sasa, from Guatemala (Sublette & Sasa, 1994 ) (new record: Costa Rica; J.H. Epler), and P. smithae (Sublette) from Mexico (see Roback, 1971). Larvae live in a variety of water bodies from streams and small pools to the littoral of lakes. Parametriocnemus Goetghebuer. One named species, P. lundbeckii (Johannsen), has been recorded from Guatemala (Sublette & Sasa, 1994). The genus has also been recorded from Costa Rica (Watson & Heyn, 1993) and southern Mex-
rectly, but tentatively assigned females from Surinam and Panama are in the ZSM collection. Nandeva Wiedenbrug, Reiss & Fittkau. This genus is known only from the Neotropical Region and Australia. Unassociated adult males from Panama (ZSM collection) cannot be confirmed as conspecific with any named species. See the descriptions and keys in Wiedenbrug et al. (1998), and Sther & Roque (2004). Recent placement of the genus in Tanytarsini (by Sther & Roque, 2004) is considered as preliminary by some workers, as the larvae are unknown. Nanocladius Kieffer. One named species is recorded: N. bubrachiatus Epler from Belize and Honduras (Spies & Reiss, 1996). It belongs to the subgenus N. (Plecopteracoluthus) Steffan in which the larvae live symphoretically or as parasites on larvae of larger aquatic insects. The genus has been recorded from Costa Rica (Watson & Heyn, 1993) and southern Mexico (Contreras Ramos & Andersen, 1999). See also Sther (1977a, which includes keys) and Epler (1986b). Larvae in the subgenus N. (Nanocladius) live freely in streams, rivers, and lakes. Neelamia Soponis. Unnamed adults are recorded from Costa Rica (Watson & Heyn, 1993). See Soponis (1987). Larvae are unknown. Nilotanypus Kieffer. Larvae of this genus have been collected in Costa Rica (Watson & Heyn, 1993), and it is also reported as unnamed adults from southern Mexico (Contreras Ramos & Andersen, 1999). See Roback (1971, 1986b). Larvae live on sandy substrates in clean, oxygen-rich running waters. Nilothauma Kieffer. Two Neotropical species originally described in this genus (Roback, 1960) are misplaced, but new generic combinations have not been established (see footnote 5 in the Key). This concerns the records of N. duena Roback and several unnamed species from Costa Rica (Watson & Heyn, 1993), as well as a male similar to N. duena from Nicaragua (ZSM collection). On the other hand, unnamed but true Nilothauma species also have been recorded from Costa Rica (Watson & Heyn, 1993). Larvae are found in lentic and lotic sediments. Oliveiriella Wiedenbrug & Fittkau. This is a new Central American record based on a rearing (J.H. Epler), including the previously unknown larva, of an undescribed species from a river in Costa Rica. See Wiedenbrug & Fittkau (1997). Adult males and larvae in the genus are indistinguishable from some Cricotopus. A comprehensive revision of all life stages of similar taxa is required to show whether generic separation is warranted. Omisus Townes. There is a single record of unnamed material of this genus from southern Mexico (Contreras Ramos & Andersen, 1999). A key to species is given in Caldwell (2000). Larvae are found in various small bodies of water, marshes and peat bogs.
474 ico (Contreras Ramos & Andersen, 1999). Larvae are known from springs and running waters. Paranilothauma Soponis. One of the two known species, P. strebulosum Adam & Sther, has been described from Costa Rica (Adam & Sther, 2000). The genus name has neuter gender, the original spelling of the species name, strebulosa, is incorrect. See also Soponis (1987). Larvae and habitat are unknown. Paraphaenocladius Thienemann. One named species has been recorded, P. exagitans Johannsen from Costa Rica and Guatemala (Sublette & Sasa, 1994; Sther & Wang, 1995). The latter authors suggest that all South American records of the genus likely belong to the subspecies P. e. longipes Sther & Wang. Larval habitats vary from terrestrial to aquatic (lentic and lotic). Paratanytarsus Thienemann & Bause. There is a single record of unnamed material of this genus from southern Mexico (Andersen et al., 2000). The globally distributed P. grimmii (Schneider) can also be expected to occur in Central America (see Langton et al., 1988). This is a parthenogenetic species with a propensity to infest water supply systems. Larvae of Paratanytarsus occur in a great variety of aquatic habitats, including brackish water. Paratendipes Kieffer. Various unnamed material is known from several Central American countries (Spies & Reiss, 1996; Contreras Ramos & Andersen, 1999). Larvae are often found in sandy sediments, some of them in hot springs (Costa Rica; Fittkau, 1999). See also Nilothauma and footnote 5 in the Key. Paratrichocladius Santos Abreu. This genus has been recorded only as larvae from Costa Rica (Watson & Heyn, 1993). The immature stages are found in various habitats, including brackish water. Parochlus Enderlein. Adults of one or more unnamed species in the P. araucanus group have been taken at highelevation sites on volcanoes in Costa Rica (Watson & Heyn, 1993; A. Borkent, unpublished). See Brundin (1966), which includes keys. Larvae of most species live in mountain streams, but some prefer springs or mosses in various small bodies of water. Pentaneura Philippi. Sublette & Sasas (1994) inclusion of Hudsonimyia Roback (see separate paragraph above) as a subgenus is not followed here. Unnamed Pentaneura material is known from Guatemala (Sublette & Sasa, 1994) and Costa Rica (Watson & Heyn, 1993). See also Roback (1971). Larvae live in various types of lotic waters. Phaenopsectra Kieffer. Pupal exuviae from Costa Rica (C. de la Rosa & W.P. Coffman, personal communication) constitute the first record of this genus from Central Amer-
ica. Phaenopsectra larvae have been found in all kinds of water bodies. Polypedilum Kieffer. This is one of the most widely distributed, common and species-rich chironomid genera. Several subgenera are recognized: Asheum Sublette & Sublette, Cerobregma Sther & Sundal, Pentapedilum Kieffer, Polypedilum Kieffer, Tripodura Townes, and Uresipedilum Oyewo & Sther (see Sther & Sundal, 1999, which includes keys to subgenera). Eleven named species are known from Central America, but about 70 from the entire Neotropical Region; see Sublette & Sasa (1994), Bidawid & Fittkau (1996), Bidawid-Kafka (1996), Spies & Reiss (1996). Larvae are found, often abundantly, in a variety of lentic and lotic habitats, including hot springs (Costa Rica; Fittkau, 1999). Chirocladius pedipalpus Picado, whose larvae live in phytotelmata (Picado, 1913), is here considered to belong in Polypedilum. Pontomyia Edwards. Species of this genus are exclusively marine. The single Central American record is from benthic samples, down to 30 m depth, around a reef island off the coast of Belize (Bretschko, 1981). Only immature stages and adult females were found. The latter are wormlike, wingless and with strongly reduced legs. Some Pontomyia species may be parthenogenetic, but adults live only for about an hour after the highly synchronised emergence, thus are easily missed in sampling. Males skate across the sea surface, propelled by their modified wings. Schrer & Epler (2007) have found larvae among algae on the backs of sea turtles caught 40 miles off a Caribbean coast. Procladius Skuse. This is a widespread, species-rich and diverse genus (e.g., see Roback, 1971). All three subgenera are likely to occur in Central America. The subgenus P. (Holotanypus) Roback is most common in the Holarctic Region, but a specimen has been found in a collection from Guatemala, although its condition does not permit species determination (C.N. Watson, unpublished). The species P. (Procladius) mozambique Roback was described from Colombia (Roback, 1982a) and is likely to occur in Central America. The subgenus P. (Psilotanypus) Kieffer is represented at least by P. (Ps.) bellus (Loew), a species widely distributed in North America and present in a collection from Panama (C.N. Watson, unpublished). Procladius (Ps.) etatus, known from Costa Rica (Watson & Heyn, 1993), may belong in the genus Laurotanypus (see above). Procladius larvae live in sediments in standing and slowly flowing waters, and many species are tolerant of low dissolved oxygen or elevated pollutant concentrations. Pseudochironomus Malloch. Unnamed specimens of various life stages have been recorded from Costa Rica (Watson & Heyn, 1993: 11 species), southern Mexico (Contreras Ramos & Andersen, 1999), and Nicaragua (Palomki, 1987). However, see footnote 3 in the Key. Larvae predominantly occur in sandy or gravelly sediments in lakes or slow-flowing rivers.
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475 build transportable cases and are found in lentic and lotic habitats. Stempellinella Brundin. Pupal exuviae from Costa Rica (C. de la Rosa & W.P. Coffman, personal communication) constitute the first record of this genus from Central America. Larvae build transportable sand cases and are found in lentic and lotic habitats. Separation of this genus from Zavrelia is unclear when all life stages are considered. Stenochironomus Kieffer. This genus is widespread and species rich, with seven named species recorded from at least one Central American country (Borkent, 1984), and more species known but undescribed (Contreras Ramos & Andersen, 1999; A. Borkent, personal communication). Larvae are miners in immersed dead wood or leaves in both lotic and lentic habitats. Stictocladius Edwards. This genus, known from the Southern Hemisphere only, requires revision to define its limits and relations. Unnamed material from Costa Rica (ZSM collection) may belong here. Larvae probably prefer relatively cool stream habitats. Sublettea Roback. Pupal exuviae from Costa Rica (C. de la Rosa & W.P. Coffman, personal communication) constitute the first record of this genus from Central America. Larvae are found in lotic habitats. Symbiocladius Kieffer. This genus shows worldwide amphitropical distribution; there are no verified records from any latitude lower than 30 degrees. Watson & Heyn (1993) reported larvae from Costa Rica, but the specimens are missing. Northern and Southern Hemisphere species separate into two subgenera with three named species each, including two each in North and South America (Caldwell, 1984; Gonser & Spies, 1997). Larvae are obligate parasites on mayfly nymphs with dorsoventrally flattened bodies, and are apparently limited to lotic habitats rarely exceeding temperatures of 20C. Synorthocladius Thienemann. This genus is recorded based on larvae from Costa Rica (Watson & Heyn, 1993). Habitats vary from lotic to lentic waters and springs. Tanypus Meigen. This is the only genus in its tribe, but it is widespread and species rich, with two recognized subgenera. Three named species have been recorded in Central America (Spies & Reiss, 1996), and more are likely to be found. See Roback (1964, 1971). The distinction between T. neopunctipennis Sublette, 1964, recorded from the eastern Nearctic Region and southern Mexico (Roback, 1971), and the western Nearctic T. grodhausi Sublette, 1964 is unclear, as rearings from southern California key to one or the other depending on the life stage (Spies, unpublished). Tanypus larvae live in sediments in standing and slowly flowing waters.
Pseudosmittia Edwards. This is a widespread and species-rich genus, but only one named species, P. forcipata (Goetghebuer), has been recorded from Central America (Sublette & Sasa, 1994). The genus has also been reported from southern Mexico (Contreras Ramos & Andersen, 1999). The larvae of most species live in terrestrial habitats, but aquatic representatives are known as well. Rheocricotopus Thienemann & Harnisch. This is the first record from Central America for this widespread genus, based on larvae from Costa Rica in the J.H. Epler collection. Holarctic species are keyed in Sther (1985). Larvae live in running waters, rarely in lake littoral zones. Rheotanytarsus Thienemann & Bause. This genus is widespread, common and rich in species. Recent reviews (Kyerematen et al., 2000; Kyerematen & Andersen, 2002) have recorded nine named species from Costa Rica, Guatemala, or Mexico. The larvae live in flowing water, where they filter-feed with silk strands spun between projections from the characteristic cases they attach to various substrates, including on the bodies of mobile animals ranging from invertebrates to catfish and alligators. Robackia Saether. This genus has not yet been found in Central America, but is known from the southern USA, and as pupae from Brazil. The Holarctic diagnosis for the adult male does not provide clear separation from all species of Parachironomus. Larvae are found in sandy sediments, most often in rivers and streams. Saetheria Jackson. Pupal exuviae from Costa Rica (C. de la Rosa & W.P. Coffman, personal communication) constitute the first record from Central America. See Jackson (1977). The current separation of this genus from Paracladopelma may not hold up if material from outside of the Holarctic Region is revised. Larvae occur in sandy sediments in lentic and lotic conditions. Skutzia Reiss. In Central America, this genus has been recorded as unnamed adults from southern Mexico (Contreras Ramos & Andersen, 1999). See Reiss (1985) and Andersen (2000). The larvae and habitat are unknown. Smittia Holmgren. There is only a single published record of this genus from the region studied here (Mexico, southern Baja California Sur; Cheng & Hogue, 1974). The small, blackish adults attract general attention only when forming mating swarms, and the larval habitats of most species, damp soil or similar accumulations of organic matter, are relatively rarely sampled. The genus is widely distributed and diverse; therefore, many species can be expected to occur in Central America. Stempellina Thienemann & Bause. In Central America, this genus has been recorded as unnamed adults from southern Mexico (Contreras Ramos & Andersen, 1999), and from Nicaragua (new record, in ZSM collection). Larvae
476 Tanytarsus van der Wulp. This is one of the most widely distributed, common, species-rich and diverse chironomid genera, but only four named species have been recorded from Central America (all described by Sublette & Sasa, 1994; note that T. guatemalensis Sublette & Sasa was accidentally omitted by Spies & Reiss, 1996). Much undescribed or undetermined material is known, e.g., 16 species from Costa Rica (Watson & Heyn, 1993), and many more species are expected to be discovered. See the keys and further references in Ekrem et al. (2003), and Sanseverino et al. (2003). The generic concept here includes species with larvae and pupae similar to those of Nimbocera Reiss, such as Tanytarsus pandus Sublette & Sasa (but see also Trivinho-Strixino & Sanseverino, 2003; Trivinho-Strixino & Strixino, 2003). Larvae of Tanytarsus occur in all types of fresh to saline water, and some species from the western Pacific are even marine. Telmatogeton Schiner. Most members of this worldwide genus live in marine habitats, and species adapted to freshwater are known only from Hawaii. Two named species have been recorded from Mexico, T. alaskensis Coquillett and T. latipennis Schiner, the latter only from around the offshore Revillagigedo Islands. Epler (unpublished) has collected pupae in southwestern Costa Rica. See Wirth (1949). Larvae live in the intertidal zone, often in the vicinity of freshwater or sewage inflows. Adults can and do fly (the first author has found T. alaskensis in a light trap located several miles inland), but most often scurry over rocks near the seas high-water mark. Tempisquitoneura Epler. The single known species, T. merrillorum Epler, was described from Costa Rica (Epler & de la Rosa, 1995). The larvae live symphoretically on Corydalus nymphs (Megaloptera). Thalassomya Schiner. Members of this genus live in marine habitats. Four named species are known from Central America (Spies & Reiss, 1996). See Wirth (1947, 1949) and Hashimoto (1979). Larvae live in the intertidal zone, often associated with reduced salinity levels and elevated organic matter loads, e.g., around harbors and river mouths. Thienemanniella Kieffer. Three named species of this widespread and species-rich genus are known from Costa Rica, Guatemala or Panama (Spies & Reiss, 1996). The genus also has been recorded from Mexico (Contreras Ramos & Andersen, 1999) and Nicaragua (Palomki, 1987). Larvae occur in all kinds of permanent aquatic habitats.
Thienemannimyia Fittkau. See footnote 2 in the Key. Based on adults, at least two species occur in Central America. These were found in Costa Rica and may be conspecific with the western Nearctic T. barberi (Coquillett) and T. fusciceps (Edwards) (C.N. Watson, unpublished). See Roback (1971). Larvae generally require relatively high dissolved oxygen concentrations, and most are rheophilic. Tribelos Townes. Contreras Ramos & Andersen (1999) have recorded adults of an unnamed species from southern Mexico. See Grodhaus (1987), which includes keys. Larvae are found in lakes, ponds, rivers and streams. Xenochironomus Kieffer. There is a single record of X. xenolabis Kieffer from Panama (Spies & Reiss, 1996). Adults of an unnamed species have been found in southern Mexico (Contreras Ramos & Andersen, 1999). Larvae are associated with freshwater sponges. Xestochironomus Sublette & Wirth. This genus is endemic to the New World; its distribution is centered around Central America, where there are eight named species (Borkent, 1984; Sublette & Sasa, 1994; Andersen & Kristoffersen, 1999), and more to be described (e.g., in the J.H. Epler and A. Borkent collections, personal communications). Larvae are miners in immersed wood in lotic habitats. Zavrelia Kieffer, Thienemann & Bause. This genus is not known from Central America, but from southern USA and Brazil. Larvae build transportable sand cases and are found in lentic and lotic habitats. See also the above entry on Stempellinella. Zavreliella Kieffer. Only one of the twelve named Neotropical species has been recorded from Central America, Z. longiseta Reiss, from Costa Rica and Panama. See Reiss (1990). Larvae build transportable, silken cases and are often associated with vegetation in eutrophic conditions. Zavrelimyia Fittkau. Members of this genus are common in the Holarctic Region but are in great need of revision (Epler, 2001). There is a single Central American record of larvae from Costa Rica (Spies & Reiss, 1996). See Roback (1971). Larvae live in a variety of habitats from springs and running waters to the littoral of lakes.
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CHIRONOMIDAE (NON-bItINg MIDgEs)

MANUAL OF CENtRAL AMERICAN DIPtERA VOLUME 1

CHIRONOMIDAE (NON-bItINg MIDgEs)

MANUAL OF CENtRAL AMERICAN DIPtERA VOLUME 1

CHIRONOMIDAE (NON-bItINg MIDgEs)

MANUAL OF CENtRAL AMERICAN DIPtERA VOLUME 1

Key to the adult males of the genera of Chironomidae of Central America

CHIRONOMIDAE (NON-bItINg MIDgEs)

MANUAL OF CENtRAL AMERICAN DIPtERA VOLUME 1

CHIRONOMIDAE (NON-bItINg MIDgEs)

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MANUAL OF CENtRAL AMERICAN DIPtERA VOLUME 1

CHIRONOMIDAE (NON-bItINg MIDgEs)

MANUAL OF CENtRAL AMERICAN DIPtERA VOLUME 1

CHIRONOMIDAE (NON-bItINg MIDgEs)

MANUAL OF CENtRAL AMERICAN DIPtERA VOLUME 1

CHIRONOMIDAE (NON-bItINg MIDgEs)

MANUAL OF CENtRAL AMERICAN DIPtERA VOLUME 1

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CHIRONOMIDAE (NON-bItINg MIDgEs)

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MANUAL OF CENtRAL AMERICAN DIPtERA VOLUME 1

CHIRONOMIDAE (NON-bItINg MIDgEs)

52. 53. 54. 55. 56.

MANUAL OF CENtRAL AMERICAN DIPtERA VOLUME 1

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62. 63. 64. 65.

CHIRONOMIDAE (NON-bItINg MIDgEs)

68. 69. 70.

MANUAL OF CENtRAL AMERICAN DIPtERA VOLUME 1

CHIRONOMIDAE (NON-bItINg MIDgEs)

74. 75. 76. 77.

79. 80. 81.

MANUAL OF CENtRAL AMERICAN DIPtERA VOLUME 1

CHIRONOMIDAE (NON-bItINg MIDgEs)

94. 95. 96.

MANUAL OF CENtRAL AMERICAN DIPtERA VOLUME 1

CHIRONOMIDAE (NON-bItINg MIDgEs)

97. 98. 99.

MANUAL OF CENtRAL AMERICAN DIPtERA VOLUME 1

CHIRONOMIDAE (NON-bItINg MIDgEs)

114. Anal point absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115 Anal point present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116

MANUAL OF CENtRAL AMERICAN DIPtERA VOLUME 1

CHIRONOMIDAE (NON-bItINg MIDgEs)

MANUAL OF CENtRAL AMERICAN DIPtERA VOLUME 1

131. Calypter with at least 1 seta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132 Calypter bare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137

CHIRONOMIDAE (NON-bItINg MIDgEs)

Synopsis of the fauna

MANUAL OF CENtRAL AMERICAN DIPtERA VOLUME 1

CHIRONOMIDAE (NON-bItINg MIDgEs)

MANUAL OF CENtRAL AMERICAN DIPtERA VOLUME 1

CHIRONOMIDAE (NON-bItINg MIDgEs)

MANUAL OF CENtRAL AMERICAN DIPtERA VOLUME 1

CHIRONOMIDAE (NON-bItINg MIDgEs)

MANUAL OF CENtRAL AMERICAN DIPtERA VOLUME 1

CHIRONOMIDAE (NON-bItINg MIDgEs)

MANUAL OF CENtRAL AMERICAN DIPtERA VOLUME 1

CHIRONOMIDAE (NON-bItINg MIDgEs)

MANUAL OF CENtRAL AMERICAN DIPtERA VOLUME 1