A revised, annotated checklist of Mexican non-biting midges (Diptera, Chironomidae)

Orestes C. Bello-González; Trond Andersen; Norman Mercado-Silva

doi:10.3897/zookeys.1191.117223

Abstract

An updated checklist of Mexican non-biting midges (Chironomidae) is presented. A total of 110 species of Chironomidae are known for Mexico: 52 species in 25 genera belong to the subfamily Chironominae, 30 species in 13 genera to Orthocladiinae, 21 species in nine genera to Tanypodinae, five species in two genera to Telmatogetoninae, and two species in one genus to Diamesinae. In addition, 41 genera without identified species are listed. The highest number of species (29) is recorded from the state of Campeche, while 19 species have been found in Veracruz and 15 in Nuevo León. Few or no records exist for states in Central and Northern Mexico, or those on the Pacific coast. The type localities for 34 species are in Mexico; of these, 27 species (25% of the total number of species recorded in the country) are endemic. Twenty-nine species recorded in Mexico have a Neotropical distribution, 15 a Nearctic distribution, and 39 species are distributed in both the Neotropical and Nearctic regions or more widely. It has been suggested that as many as 1000 species might occur in Mexico; so only a little more than 10% of the expected diversity has so far been recorded.

Key words

Biodiversity, Nearctic, Neotropical, transition zone

Introduction

Mexico is a megadiverse country (Mittermeier et al. 2011; Mendoza-Ponce et al. 2020). Located in the Nearctic-Neotropical transition area, the north to south orientation of numerous warm, low altitude corridors, and the abundance of mountain chains with colder conditions have allowed biota to disperse during past climate change events (Halffter 1987). This high biodiversity results primarily from an accumulation of taxa from other areas and constant changes in the landscape (Priego Santander and Esteve Selma 2017) rather than local diversification (Sundaram et al. 2019; Harvey et al. 2020). However, several of the most species rich and ecologically relevant insect groups are not included in these studies.

Chironomidae have the widest distribution of all free-living groups of holometabolous insects and are likely the most taxonomically and ecologically diverse family of aquatic insects (Cranston 1995; Dijkstra et al. 2014). Reiss (1982) estimated that between 1500–2000 chironomid species might occur in tropical Mexico and Central America, and Andersen et al. (2000) suggested that as many as 1000 species can be expected to occur in Mexico. Spies et al. (2009) compiled an annotated list of the Mexican and Central American genera including a key to the genera known from the region at that time. However, the latest inventory of the Mexican chironomids only included 61 species plus an additional 25 genera without identified species (Andersen et al. 2000).

During the last two decades, several new species have been described based on material from Mexico (Kyerematen et al. 2000; Kyerematen and Andersen 2002; Wang et al. 2006; Vinogradova 2008; Andersen et al. 2010, 2016; Wiedenbrug et al. 2012; Pinho et al. 2013; Acosta et al. 2017; Pinho and Andersen 2021; Andersen 2023). Several species and genera have also been recorded for the first time from Mexico, especially in connection with surveys of particular habitats like the aquatic fauna in spring-fed tropical canyons in the southern Sonora desert (Bogan et al. 2014), or subfossil Chironomidae in surface sediments of the sinkholes of the Yucatan Peninsula (Hamerlík et al. 2018). Several of these studies are based mainly on larvae and the materials are generally not identified beyond genus level.

The Nearctic-Neotropic transition should lead to the existence of chironomid species with different biogeographic affinities. The Nearctic fauna is comparatively well known (Oliver et al. 1990; Oliver and Dillon 1994) and most chironomids in Mexico with this biogeographic affinity can be identified to genus level using the keys to the larvae, pupa and adults of the Holarctic Region (Wiederholm 1986, 1989; Andersen et al. 2013a). The Neotropical chironomids from Mexico are much less studied and more difficult to identify based on available literature.

An updated checklist of Mexican Chironomidae species is presented. The list provides an updated baseline and will facilitate the study of the chironomid fauna in the Nearctic-Neotropical biogeographical transition zone in Mexico. The checklist is based on Andersen et al. (2000), and new records and species published during the last two decades are added. Some ecological information now available for the genera recorded from Mexico are also included.

Methods

The checklist is based on Andersen et al. (2000); references already given in that list are not repeated here. The checklist includes published records only. Records were compiled from peer reviewed scientific articles, books, and book chapters and, to a lesser extent, unpublished project reports. Specimens of Mexican chironomids are housed in several collections (Contreras-Ramos 2021; Huerta Jiménez 2021; Admin 2022; Bentley and Thomas 2022; European Bioinformatics Institute 2022); and these records can be accessed using “Name search” in GBIF (2023).

Following Ashe and O´Connor (2009) eight major zoogeographical regions are recognized: Antarctic (AN), Neotropical (NT), Nearctic (NE), Palaearctic (PA), Afrotropical (AF), Oriental (OR), Australasian (AU), and Oceanian (OC). Administratively, Mexico is divided in 32 states. Of these the 18 northernmost states are generally regarded as belonging to the Nearctic region, while the remaining 13 southern most states as belonging to the Neotropical region (Ashe and O´Connor 2009). However, the biogeographical zones are not clearly defined and depend to some degree on the group of organisms studied. There are also clearly transition zones between the two regions. Given this, taxa present in Mexico and in Central or South America are considered to be Neotropical, while taxa present in Mexico and in the USA and/or Canada are considered to be Nearctic. The exception is taxa from southern Florida, USA, which is considered to be Neotropical. However, many species are found both in South- and North America or have a wider distribution.

The checklist is arranged alphabetically. Species group names follow the genus and subfamily names. A short outline with information on the number of species, distribution, and larvae habitats is given for each genus. For literature records given as “Cricotopus cf. sylvestris” or “Cricotopus sylvestris group” we assume they are correctly identified to genus level. Following Ashe and O’Connor (2009), two Tanypodinae species originally described as Macropelopia roblesi Vargas, 1946 and Pentaneura marmorata Johannsen, 1938 are listed as “Generically unplaced valid Macropelopiini” and “Generically unplaced valid Tanypodinae”, respectively.

The valid species name is followed by the original combination in parenthesis, with type country (for USA, country and state) in square brackets. When the type locality is situated in Mexico, more specific information is given for the type locality. Synonyms are given if descriptions are based on Mexican material. Mexican records are then given followed by the state(s) from where the species is recorded in square brackets. Finally, the total distribution for a species is given as zoogeographical region(s), followed by the countries from where the species has been recorded. However, this list of countries might not be complete, and for widespread species it is only given as “widespread”.

Citations for species names are arranged as follows

Cladopelma forcipis (Rempel, 1939: 211) (Chironomus (Cryptochironomus)) [Brazil]. Andersen et al. (2000: 590) [Mexico State; Morelos; Veracruz]; Vinogradova and Riss (2007: 34) [“Yucatan Peninsula”]; European Bioinformatics Institute (2022) [Quintana Roo]. NT, NE. Brazil, Colombia, Costa Rica, Guatemala, Mexico, Nicaragua, Panama, USA.

In the checklist, recorded genera lacking identified species are included. Mexican records are listed as e.g.: Cryptochironomus sp., followed by the state(s) from where the genus is recorded in square brackets.

Vinogradova and Riss (2007) provided numerous records from the Yucatan Peninsula, but without giving any details on the localities. As Yucatan Peninsula includes partly or totally the territory of three Mexican states; these records are listed as [“Yucatan Peninsula”].

Results

Check list

Subfamily Chironominae

Genus Apedilum Townes, 1945

A genus with three named species. A. elachistus Townes, 1945 is widespread throughout North and South America, A. subcinctum Townes, 1945 is distributed in North and Central America, and A. griseistriatum (Edwards, 1931) occurs in South America. Larvae are associated with submerged vegetation in ponds, canals, lakes, and slowly running rivers, both in fresh or brackish water (Epler et al. 2013).

Apedilum elachistus Townes, 1945: 33 [USA: Texas]. Andersen et al. (2000: 590) [States of Campeche; Puebla; Veracruz]; Contreras-Ramos et al. (2000: 25); Vinogradova and Riss (2007: 33) [“Yucatan Peninsula”]. NT, NE. Argentina (Donato et al. 2008a), Brazil, Canada (Giberson et al. 2001), Costa Rica, Guatemala, Mexico, Nicaragua, Uruguay (Donato et al. 2008a), USA.

Apedilum subcinctum Townes, 1945: 33 [USA: Nevada]. Andersen et al. (2000: 590) [States of Campeche; Jalisco]; Contreras-Ramos et al. (2000: 25); Vinogradova and Riss (2007: 33) [“Yucatan Peninsula”]; Contreras-Ramos (2021). NT, NE. Guatemala, Mexico, USA.

Genus Asheum Sublette & Sublette, 1983

See: Polypedilum Kieffer, 1912.

Genus Axarus Roback, 1980

A genus of ~ 15 species that occur in the Neotropical, Nearctic, Palaearctic, and the Australasian regions. Ten species are known from South America (Andersen and Mendes 2002a; Andersen et al. 2018; Pinho et al. 2019). Larvae occur in littoral to sublittoral soft sediments in lakes and rivers (Epler et al. 2013).

Axarus rogersi (Beck & Beck, 1958: 27) (Xenochironomus) [USA: Florida]. Andersen et al. (2000: 590) [Campeche State]; Contreras-Ramos et al. (2000: 26); Contreras-Ramos (2021). NT, NE. Costa Rica, Mexico, Nicaragua, USA.

Genus Beardius Reiss & Sublette, 1985

A genus with > 30 named species that occur mainly in tropical areas in the Neotropical region with a few species in the southern parts of the Nearctic region (Jacobsen and Perry 2000; Pinho et al. 2013). The larvae have been found associated with macrophytes or submerged wood in both standing and flowing waters (Epler et al. 2013).

Beardius aciculatus Andersen & Sæther, 1996: 40 [Costa Rica]. Andersen et al. (2000: 590) [States of Campeche; Veracruz]; Contreras-Ramos et al. (2000: 26); Vinogradova and Riss (2007: 33) [“Yucatan Peninsula”]; Contreras-Ramos (2021); Admin (2022). NT. Costa Rica, Mexico.

Beardius chapala Pinho, Mendes & Andersen, 2013: 28 [Mexico: Jalisco State, Lake Chapala, El Chante]. Endemic.

Beardius parcus Reiss & Sublette, 1985: 183 [Venezuela]. Andersen et al. (2000: 590) [Veracruz State]; Contreras-Ramos et al. (2000: 26) [Campeche State]; Contreras-Ramos (2021). NT. Costa Rica, Mexico, Nicaragua, Venezuela.

Genus Caladomyia Säwedal, 1981

See: Tanytarsus Wulp, 1874.

Genus Chironomus Meigen, 1803

One of the most species-rich and common chironomid genera, with ~ 300 described species from all zoogeographical regions except Antarctica. The larvae graze on detritus or are filter-feeders, predominantly in soft sediments of standing water, rarely in flowing water (Epler et al. 2013).

Chironomus alchichica Acosta & Prat in Acosta et al. 2017: 53 [Mexico: Puebla State, Lake Alchichica]. Endemic.

Chironomus stigmaterus Say, 1823: 15 [USA: Pennsylvania]. Andersen et al. (2000: 590) [States of Durango; Puebla]; Alcocer et al. (2016: 411). NT, NE. Brazil, Cuba, Mexico, USA.

Genus Cladopelma Kieffer, 1921

A genus of ~ 20 described species that occur in all zoogeographical regions except Antarctica and Oceania. The larvae live in streams and larger rivers, lakes, and ponds as well as brackish water and hot springs (Epler et al. 2013).

Cladopelma forcipis (Rempel, 1939: 211) (Chironomus (Cryptochironomus)) [Brazil]. Andersen et al. (2000: 590) [States of Mexico; Morelos; Veracruz]; Vinogradova and Riss (2007: 34) [“Yucatan Peninsula”]; European Bioinformatics Institute (2022) [Quintana Roo State]. NT, NE. Brazil, Colombia, Costa Rica, Guatemala, Mexico, Nicaragua, Panama, USA.

Genus Cladotanytarsus Kieffer, 1921

A genus of ~ 80 described species that occur in all zoogeographical regions except Antarctica and Oceania. No named species are recorded from South America, but larval morphotypes have been recorded from Brazil (Roque et al. 2004). Larvae construct sessile cases of fine detritus and have been found in streams and larger rivers, lakes, and ponds, as well as in brackish water and hot springs (Epler et al. 2013).

Cladotanytarsus viridiventris (Malloch, 1915: 491) (Tanytarsus) [USA: Michigan]. Andersen et al. (2000: 590) [Puebla State]. NE. Canada, Mexico, USA.

Genus Cryptochironomus Kieffer, 1918

A genus of ~ 60 named species that occur in all zoogeographical regions, except Antarctica. Four species are described from South America (da Silva et al. 2010). Larvae occur on various substrates in lakes, small streams, and larger rivers (Epler et al. 2013).

Cryptochironomus sp.: Vinogradova and Riss (2007: 34) [“Yucatan Peninsula”]; Granados-Ramírez et al. (2017: 45) [Morelos State].

Genus Dicrotendipes Kieffer, 1913

A genus of ~ 85 described species that occur in all zoogeographic regions except Antarctica. The genus was revised by Epler (1988). The larvae inhabit the littoral sediments of standing waters and may be common in lentic habitats (Epler et al. 2013).

Dicrotendipes aethiops (Townes, 1945: 107) (Tendipes (Limnochironomus)) [USA: New Mexico].

Syn.: Tendipes (Limnochironomus) figueroai Vargas, 1952: 48 [Mexico: Morelos State].

Andersen et al. (2000: 590) [States of Baja California; Mexico; Querétaro]; Huerta-Jiménez (2021) [Morelos State]. NE. Mexico, USA.

Dicrotendipes californicus (Johannsen, 1905: 217) (Chironomus) [USA: California]. Andersen et al. (2000: 590) [States of Mexico; Morelos; Oaxaca; Sinaloa]; Bentley and Thomas (2022) [Michoacán State]. NT, NE. Chile, Colombia, Costa Rica, Guatemala, Mexico, Panama, Peru, USA.

Remark. Bentley and Thomas (2022) recorded the species from “Cojumatlán, Jalisco”. The town of Cojumatlán is located on the shoreline of Lake Chapala in the State of Michoacán. Although close to the border between the two states, the original reference to the State of Jalisco most probably is a mistake.

Dicrotendipes neomodestus (Malloch, 1915: 475) (Chironomus) [USA: Illinois]. Andersen et al. (2000: 590) [Puebla State]; Alcocer et al. (2016: 412). NE. Canada, Mexico, USA.

Dicrotendipes obrienorum Epler, 1987: 148 [Mexico: Michoacán State, Patzcuaro]. Andersen et al. (2000: 590). Endemic.

Dicrotendipes sinoposus Epler, 1987: 152 [Mexico: Hidalgo State, Otongo]. Andersen et al. (2000: 590) [States of Campeche; Hidalgo; Veracruz]; Contreras-Ramos et al. (2000: 26); Vinogradova and Riss (2007: 34) [“Yucatan Peninsula”]; Contreras-Ramos (2021). NT. Brazil, Colombia, Costa Rica, Dominica, Mexico, Nicaragua.

Genus Einfeldia Kieffer, 1922

The concept, content and status of Einfeldia have been, and to an extent remain, confusing (Cranston et al. 2016a). Narrowly defined, Einfeldia contains approximately five species and is distributed in the Neotropical, Nearctic, Palaearctic, Oriental, and Australasian regions. The larvae inhabit standing, predominantly dystrophic waters (Epler et al. 2013).

Einfeldia sp.: Navarrete-Salgado et al. (2004: 157) [México State]; Vinogradova and Riss (2007: 34) [“Yucatan Peninsula”].

Genus Endochironomus Kieffer, 1918

A genus of ~ 20 named species distributed in the Nearctic, Palaearctic, Afrotropical, and Oriental regions. The larvae occur in “Aufwuchs” on living and dead substrata in almost all types of still water; they mine leaves and stems of macrophytes preferentially in small, eutrophic standing waters (Epler et al. 2013).

Endochironomus subtendens (Townes, 1945: 65) (Tanytarsus (Endochironomus)) [USA: New York]. Andersen et al. (2000: 590) [Yucatán State]. NE. Canada, Mexico, USA.

Genus Endotribelos Grodhaus, 1987

A genus of 14 described species, all occurring in the Neotropical and Nearctic regions except one species from China. The Brazilian species were treated by Trivinho-Strixino and Pepinelli (2015). The larvae are associated with aquatic macrophytes, decaying leaves, wood, and fallen fruits in streams (Epler et al. 2013).

Endotribelos hesperium (Sublette, 1960: 217) (Tendipes (Tribelos)) [USA: California]. Andersen et al. (2000: 590) [Puebla State]. NE. Mexico, USA.

Genus Fissimentum Cranston & Nolte, 1996

A genus with four named species endemic to South America; but larval morphotypes have also been recorded from the Nearctic, Oriental, and Australasian regions (Epler et al. 2013). Larvae are found in fine sediments of lentic and lotic habitats and can tolerate desiccation (Cranston and Nolte 1996).

Fissimentum sp.: Vinogradova and Riss (2007: 33) [“Yucatan Peninsula”].

Genus Glyptotendipes Kieffer, 1913

The taxonomy and nomenclature of the genus have been confusing since its establishment. Glyptotendipes now includes ~ 27 species, distributed in the Neartic, Paleartic, Oriental, and Afrotropical regions (Epler et al. 2013; Konar and Majumdar 2021). Three subgenera are recognized, Glyptotendipes s. str. (including Phytotendipes Goetghebuer, 1937), Caulochironomus Heyn, 1992, and Heynotendipes Spies & Sæther, 2004 (including Trichotendipes Heyn, 1992) (see Spies and Sæther 2004). Larvae occur in detritus-rich littoral sediments of lakes, ponds, small water bodies, and running water (Epler et al. 2013).

Glyptotendipes sp.: Contreras-Ramos and Andersen (1999: 4) [Campeche State]; Vinogradova and Riss (2007: 34) [“Yucatan Peninsula”].

Genus Goeldichironomus Fittkau, 1965

A genus of 15 named species mainly distributed in the Neotropical region (tropical and subtropical Central and South America), but several species reach their northern limits in southeastern USA (Donato and Andersen 2022). The larvae of Goeldichironomus are mostly found in sediments, on plants or in floating mats of vegetation in lentic habitats, in fresh to brackish water, and in oligotrophic to hypereutrophic conditions (Epler et al. 2013).

Goeldichironomus amazonicus (Fittkau, 1968: 260) (Siolimyia) [Brazil]. Andersen et al. (2000: 590) [Veracruz]; Vinogradova and Riss (2007: 34) [“Yucatan Peninsula”]. NT. Bahamas, Brazil, Mexico, Nicaragua, Panama, Peru, USA: Florida, Venezuela, Virgin Islands.

Goeldichironomus carus (Townes, 1945: 118) (Tendipes) [Venezuela]. Vinogradova and Riss (2007: 32) [“Yucatan Peninsula”]. NT, NE. Colombia, Costa Rica, Mexico, Nicaragua, Panama, Venezuela, USA.

Goeldichironomus holoprasinus (Goeldi, 1905: 135) (Chironomus) [Brazil]. Andersen et al. (2000: 590) [Tabasco]; Vinogradova and Riss (2007: 34) [“Yucatan Peninsula”]. NT, NE. Argentina, Brazil, Costa Rica, Ecuador, Mexico, Nicaragua, Panama, Peru, USA, Venezuela, Virgin Islands.

Genus Harnischia Kieffer, 1921

A genus of ~ 20 described species that occur in all zoogeographic regions except the Antarctic, Neotropical, and Oceanian regions. However, unnamed larvae have been recorded from Brazil (Roque et al. 2004). Larvae occur in soft sediments of generally clean lakes and larger rivers (Epler et al. 2013).

Harnischia sp.: Contreras-Ramos et al. (2000: 26) [Campeche State].

Genus Hyporhygma Reiss, 1982

A genus with a single named species, H. quadripunctatum (Malloch, 1915), distributed in eastern North America, from Newfoundland to Florida. The larvae mine leaves and stems of Nuphar and Nymphaea species (Epler et al. 2013).

Hyporhygma sp.: Vinogradova and Riss (2007: 34) [“Yucatan Peninsula”].

Genus Kiefferulus Goetghebuer, 1922

Syn.: Nilodorum Kieffer, 1921 (see Cranston et al. 1990).

A genus with at least five species in the Holarctic region. Species previously considered to belong to Nilodorum are widespread in the Afrotropical, Oriental, and Australasian regions. The larvae inhabit sediments of small to medium sized waterbodies (Epler et al. 2013).

Kiefferulus sp.: Contreras-Ramos et al. (2000: 26) [Campeche State].

Genus Lauterborniella Thienemann & Bause, 1913

A genus with a single named species, L. agrayloides (Kieffer, 1911) distributed in the Neotropical, Nearctic, and Palaearctic regions. Other species referred to as Lauterborniella in the literature belong either to Zavreliella Kieffer, 1920 or to Kribiodorum Kieffer, 1921, or their generic affinities are unclear (Epler et al. 2013). Larvae are mobile amongst submerged vegetation in small bodies of standing water (Epler et al. 2013). In Brazilian streams they have also been found in accumulations of litter attached to stones (Sanseverino and Nessimian 2001).

Lauterborniella sp.: Bogan et al. (2014: 2726) [Sonora State].

Genus Microchironomus Kieffer, 1918

A genus of approximately ten species distributed in the Nearctic, Palaearctic, Afrotropical, and Oriental regions (Yan and Wang 2006). The larvae occur in lakes, rivers, and ditches, including brackish water (Epler et al. 2013).

Microchironomus nigrovittatus (Malloch, 1915: 456) (Chironomus) [USA: Illinois]. Andersen et al. (2000: 590) [Veracruz State]. NE. Mexico, USA.

Genus Micropsectra Kieffer, 1909

Based on morphological and molecular data, Krenopsectra Reiss, 1969 and Parapsectra Reiss, 1969 were recently considered to be junior synonyms of Micropsectra (Ekrem et al. 2010). The three genera have a Holarctic distribution with ~ 100 valid species. The larvae have been recorded from a wide range of habitats, including hygropetric situations, thermal springs, moorland pools, and temporary pools. They are often found in muddy deposits in slow flowing stretches of streams and small rivers and in mesotrophic and oligotrophic lakes (Epler et al. 2013).

Micropsectra sp.: Vinogradova and Riss (2007: 34) [“Yucatan Peninsula”]; Alcocer et al. (2016: 411) [Puebla State].

Remarks. Both records from Mexico appear to be based on larvae only. According to Epler et al. (2013) the larvae of Micropsectra can be difficult to separate from Tanytarsus larvae.

Genus Microtendipes Kieffer, 1915

A genus of ~ 55 named species that occur in all zoogeographic regions, except Antarctica; Neotropical records are based only on larvae not identified to species level (Roque et al. 2004). Larvae are found in littoral and sublittoral sediments of lakes, and in sediments and submerged mosses in running water (Epler et al. 2013).

Microtendipes sp.: Vinogradova and Riss (2007: 34) [“Yucatan Peninsula”]; Granados-Ramírez et al. (2017: 45) [Mexico State].

Genus Nandeva Wiedenbrug, Reiss & Fittkau, 1998

A genus with seven described species that occur in the Neotropical and Australasian regions (Andersen et al. 2011b). The only described larvae was found in semi-immersed leaf litter packs in a tropical stream in Australia (Cranston 2019).

Nandeva strixinorum Sæther & Roque, 2004: 67 [Brazil]. Andersen et al. (2011b: 55) [Campeche State]. NT. Brazil, Mexico.

Genus Nilothauma Kieffer, 1921

A genus with > 60 described species distributed throughout most zoogeographic regions except Antarctica. The Neotropical species were reviewed by Pinho and Andersen (2021). The larvae inhabit littoral and sublittoral soft sediments of lakes, streams, and rivers (Epler et al. 2013).

Nilothauma maya Pinho & Andersen, 2021: 103 [Mexico: Campeche State, Calakmul]. Endemic to Mexico.

Genus Nimbocera Reiss, 1972

See: Tanytarsus Wulp, 1874.

Genus Omisus Townes, 1945

See: Zavreliella longiseta Reiss, 1990.

Genus Oukuriella Epler, 1986

A genus of > 20 species restricted to the Neotropical region. The larvae can be found associated with freshwater sponges or submerged wood in streams and rivers (Fusari et al. 2014). Species associated with sponges were revised by Fusari et al. (2014).

Oukuriella annamae Epler, 1996: 4 [Costa Rica]. Andersen et al. (2000: 590) [Campeche State]; Contreras-Ramos et al. (2000: 26). NT. Brazil (Bellodi et al. 2016), Costa Rica, Mexico.

Oukuriella oliveirai Messias & Fittkau, 1997: 256 [Brazil]. Bellodi et al. (2016: 191) [Campeche State]. NT. Brazil, Mexico.

Oukuriella simulatrix Epler, 1986: 160 [Colombia]. Andersen et al. (2000: 590) [Campeche State]; Contreras-Ramos et al. (2000: 26); Vinogradova and Riss (2007: 33) [“Yucatan Peninsula”]; Bellodi et al. (2016: 193). NT. Colombia, Mexico.

Genus Parachironomus Lenz, 1921

The genus has a worldwide distribution with at least 30 species in the Holarctic region and 20 species in the Neotropical region (Trivinho-Strixino et al. 2010; Epler et al. 2013). The adults of the Neotropical species were revised by Spies et al. (1994). Larvae are found in lentic and lotic water bodies under a wide range of conditions, including leaf miners in submerged macrophytes; they also live in association with Bryozoa or are ectoparasites on other invertebrates (Epler et al. 2013).

Parachironomus directus (Dendy & Sublette, 1959: 514) (Tendipes (Cryptochironomus)) [USA: Alabama]. Andersen et al. (2000: 590) [Morelos State]; Vinogradova and Riss (2007: 33) [“Yucatan Peninsula”]. NT, NE. Mexico, Nicaragua, Panama, USA.

Parachironomus hazelriggi Spies, 2000: 133 [USA: California]. Andersen et al. (2000: 590) as P. monochromus Wulp, 1874 [States of Querétaro; Mexico]; Spies (2000: 134) [Guanajuato State; Mexico City]. NE, PA. Canada, Mexico, Russia (Orel 2017: 535), USA.

Remarks. According to Spies (2000: 129) P. monochromus Wulp, 1874 is today considered to be a Palaearctic species and listing from Mexico following Spies & Reiss (1996: 71) must be changed to P. hazelriggi.

Parachironomus tenuicaudatus (Malloch, 1915: 475) (Chironomus) [USA: Illinois]. Andersen et al. (2000: 590) [Puebla State]. NE, PA. Widespread.

Remarks. According to Spies (2000: 133) the record from Puebla is based on Alcocer et al. (1993) and must be considered as uncertain as it appears to be based on immature specimens only.

Genus Paracladopelma Harnisch, 1923

The genus has a predominantly Holarctic distribution, with at least 20 known species; many species are also recorded from the Oriental region (Epler et al. 2013; Yan et al. 2008). The Holartic species were reviewed by Jackson (1977). Larvae inhabit sandy substrata in lakes, streams, and small rivers and the soft profundal sediments of deep lakes (Epler et al. 2013).

Paracladopelma sp.: Vinogradova and Riss (2007: 33) [“Yucatan Peninsula”].

Genus Paralauterborniella Lenz, 1941

A genus with two described species; one of them, P. nigrohalteralis (Malloch, 1915), is widely distributed (Tang 2016). The larvae usually occur in littoral soft sediments of lakes (Epler et al. 2013).

Paralauterborniella sp.: Vinogradova and Riss (2007: 33) [“Yucatan Peninsula”].

Genus Paratanytarsus Thienemann & Bause, 1913

A genus of > 60 named species that occur in all zoogeographical regions except Antarctica. The larvae inhabit brackish ponds, cool streams, lakes, rivers, reservoirs, and marshes (Epler et al. 2013).

Paratanytarsus tolucensis Reiss, 1972: 62 [Mexico: Mexico State, Nevado de Toluca]. Andersen et al. (2000: 590). Endemic.

Genus Paratendipes Kieffer, 1911

A genus of nearly 40 named species that occur in the Afrotropical and Oriental regions and in the Holarctic realm (Qi et al. 2009). For South America there are only records of unnamed species (Roque et al. 2004; Trivinho-Strixino 2011). Larvae are found in lakes, ponds, small water bodies, bogs, and hot springs and in streams and rivers in soft sediments and sandy bottoms (Epler et al. 2013).

Paratendipes sp.: Contreras-Ramos and Andersen (1999: 4) [Campeche State]; Vinogradova and Riss (2007: 33) [“Yucatan Peninsula”]; Bogan et al. (2014: 2726) [Sonora State]; Hamerlík et al. (2018: 217) [Yucatan State].

Genus Phaenopsectra Kieffer, 1921

A genus of more than ten named species that occur in all zoogeographical regions except the Antarctic, Oriental, and Australasian regions. The larvae mainly occur in sandy and muddy sediments of small standing and flowing waters, but also on submerged water plants and hard substrata (Epler et al. 2013).

Phaenopsectra sp.: Navarrete-Salgado et al. (2004: 157) [México State]; Vinogradova and Riss (2007: 33) [“Yucatan Peninsula”].

Genus Polypedilum Kieffer, 1912

Asheum Sublette & Sublette, 1983, as subgenus

Syn.: Pedionomus Sublette, 1964 (see Sæther and Sundal 1999).

The largest genus of Chironomidae, with > 500 described species that occur in all zoogeographical regions except Antarctica. Based on imaginal characters, eight subgenera were recognized by Sæther et al. (2010), namely Tripedilum Kieffer, 1921; Polypedilum s. str.; Pentapedilum Kieffer, 1913; Tripodura Townes, 1945; Uresipedilum Oyewo & Sæther, 1998; Cerobregma Sæther & Sundal, 1999; Kribionympha Kieffer, 1921; and Probolum Andersen & Sæther, 2010. However, the delimitation of the subgenera within Polypedilum was questioned by Yamamoto and Yamamoto (2015) and Cranston et al. (2016b). The position of Asheum is unclear but is usually treated as a subgenus within Polypedilum (see Pinho and Silva 2020). Larvae of Polypedilum occur in virtually all still and flowing waters, except in the Arctic and at high elevation. They are mostly found in sediments, mining water plants or specializing in plant-held waters (phytotelmata) (Epler et al. 2013).

Polypedilum (Asheum) beckae (Sublette, 1964a: 137) (Pedionomus) [USA: Louisiana]. Andersen et al. (2000: 590) [States of Campeche; Veracruz]; Contreras-Ramos et al. (2000: 25). NT, NE. Dominican Republic, Mexico, USA.

Polypedilum (Asheum) curticaudatum Rempel, 1939: 214 [Brazil]. Vinogradova and Riss (2007: 33) (as: Pedionomus curticaudatus) [“Yucatan Peninsula”]; Pinho and Silva (2020: 184). NT. Brazil, Mexico.

Polypedilum (Polypedilum) purus Bidawid-Kafka, 1996: 216 [Brazil]. Vinogradova and Riss (2007: 33) [“Yucatan Peninsula”]. NT. Brazil, Mexico.

Polypedilum (Tripodura) bacalar Vinogradova, 2008: 279 [Mexico: Quintana Roo State, Chetumal, Lake Bacalar]; Zhang et al. (2016: 41). Endemic.

Polypedilum (Tripodura) rissi Vinogradova, 2008: 281 [Mexico: Yucatan State, Lake Punta Laguna]. Vinogradova (2008: 281) [Quinto Roo]; Zhang et al. (2016: 47). NT. Guatemala, Mexico

Polypedilum (Tripodura) spiesi Vinogradova, 2008: 278 [Belize]. Vinogradova (2008: 278) [Quintana Roo State]; Zhang et al. (2016: 48). NT. Belize, Mexico.

Polypedilum (Uresipedilum) pedatum Townes, 1945: 55 [USA: New York & Washington]. Andersen et al. (2000: 590) [Nuevo León State]. NE. Canada, Mexico, USA.

Remarks. Townes (1945: 55) described two subspecies, P. pedatum pedatum from New York and P. pedatum excelsius from Washington. Andersen et al. (2000: 590) recorded the species as P. (Polypedilum) pedatum Townes, while Sæther and Oyewo (2008: 3) placed it in subgenusUresipedilum.

Polypedilum rohneri Vinogradova, 2008: 286 [Belize]. Vinogradova (2008: 286) [Yucatan State]. NT. Belize, Guatemala, Mexico.

Remarks. The species is not assigned to a subgenus. According to Vinogradova (2008: 288) it might deserve a separate subgenus.

Genus Pseudochironomus Malloch, 1915

A genus with at least 30 species distributed in the Neotropical, Nearctic, and Palaearctic regions. The Brazilian species have recently been treated by Shimabukuro et al. (2017) and Trivinho-Strixino and Shimabukuro (2018). The larvae inhabit sandy or gravelly littoral sediments, primarily in meso- or oligotrophic lakes or in large, slow flowing rivers (Epler et al. 2013).

Pseudochironomus seipi Andersen, 2023 [Mexico: Chiapas State, Chintul, Río Chintul]. NT. Costa Rica, Mexico.

Genus Rheotanytarsus Thienemann & Bause, 1913

A genus with ~ 100 species distributed in all zoogeographic regions except Antarctica. The Central American and Mexican species were reviewed by Kyerematen and Andersen (2002); the Rheotanytarsus pellucidus group was revised by Kyerematen et al. (2000). Larvae are rheobiontic, occurring in streams, large rivers, and the littoral of lakes where wave action simulates the action of flowing water (Epler et al. 2013).

Rheotanytarsus calakmulensis Kyerematen & Andersen, 2002: 33 [Mexico: Campeche State, Calakmul Biosphere Reserve]. Endemic.

Rheotanytarsus contrerasi Andersen & Sæther in Kyerematen et al., 2000: 166 [Mexico: Puebla State, Mpio. Progreso, Río San Juan]. Kyerematen et al. (2000: 166) [Nuevo León State]. Endemic.

Rheotanytarsus foliatus Kyerematen & Andersen, 2002: 35 [Costa Rica]. Kyerematen and Andersen (2002: 35) [Nuevo León State]. NT. Costa Rica, Mexico.

Rheotanytarsus hanseni Kyerematen & Andersen, 2002: 42 [Mexico: Oaxaca State, Candelaria Loxiela]. Kyerematen and Andersen (2002: 42) [Morelos State]. Endemic.

Rheotanytarsus kusii Kyerematen & Andersen, 2002: 37 [Mexico: Nuevo León State, Allende, Río Ramos]. Endemic.

Rheotanytarsus nuamae Kyerematen & Andersen, 2002: 38 [Mexico: Nuevo León State, Allende, Río Ramos]. Endemic.

Rheotanytarsus ramirezae Kyerematen & Andersen, 2002: 46 [Mexico: Nuevo León State, Santiago, Cola de Caballo]. Endemic.

Genus Saetheria Jackson, 1977

A genus of seven named species that occur in the Neotropical, Nearctic, and Palearctic regions (Orel 2014). Only unnamed larvae have so far been recorded from South America (Roque et al. 2004). Larvae inhabit sandy substrata of lakes and streams (Epler et al. 2013).

Saetheria sp.: Vinogradova and Riss (2007: 33) [“Yucatan Peninsula”].

Genus Skutzia Reiss, 1985

A genus of six species that occur in the Neotropical, Nearctic, and Oriental regions. The genus was revised by Pinho et al. (2009a). The larvae are unknown. However, they can be expected to construct transportable cases of sand grains, small wood or plant remains, as seen in the larvae of other species in the subtribe Zavreliina.

Skutzia quetzali Pinho, Mendes & Andersen, 2009a: 204 [Mexico: Campeche State, Calakmul, Ejido Nuevo Becan, El Chorro]. NT. Mexico, Panama.

Genus Stempellina Thienemann & Bause, 1913

A genus of at least 20 species that occur in all zoogeographic regions except Antarctica. The larvae construct long, curved, tapered, transportable cases of fine sand and silt. They are eurytopic, occurring in springs, streams, larger rivers, lakes, brackish water, moorland pools, and in thermal springs (Epler et al. 2013).

Stempellina sp.: Contreras-Ramos and Andersen (1999: 4) [Campeche State]; Vinogradova and Riss (2007: 34) [“Yucatan Peninsula”]; Bogan et al. (2014: 2726) [Sonora State].

Genus Stempellinella Brundin, 1947

A genus of ~ 20 described species that occur in all zoogeographical regions except Antarctica. The larvae construct straight, transportable cases of fine sand and silt, often speckled with detritus; they occur in unpolluted springs and small streams as well as in lakes (Epler et al. 2013).

Stempellinella sp.: Vinogradova and Riss (2007: 34) [“Yucatan Peninsula”]; Bogan et al. (2014: 2726) [Sonora State].

Genus Stenochironomus Kieffer, 1919

A genus of > 100 described species that occur in all zoogeographic regions except Antarctica. The genus was revised by Borkent (1984). South American species were treated by Dantas et al. (2016). Larvae are obligate miners in living or dead vegetation including woody parts of plants, in both lentic and lotic situations (Epler et al. 2013).

Stenochironomus leptopus Kieffer, 1906: 19 [St. Vincent]. Andersen et al. (2000: 590) [Mexico, without specific locality]. NT. Costa Rica, Dominica, Ecuador, Guatemala, Mexico, St. Vincent.

Genus Sublettea Roback, 1975

A small genus with four species distributed in the Neotropical, Nearctic, and Oriental regions (Ashe et al. 1987). The larvae occur in flowing waters including cool, clean, fast flowing, temperate streams and warm, tropical rivers and streams (Epler et al. 2013). The only known larva construct soft, non-transportable cases of fine granules and silk that are attached to the substrate (Roback 1975).

Sublettea sp.: Vinogradova and Riss (2007: 34) [“Yucatan Peninsula”].

Genus Tanytarsus Wulp, 1874

Syn.: Nimbocera Reiss, 1972 (see Sanseverino et al. 2010).

Syn.: Caladomyia Säwedal, 1981 (see Lin et al. 2018).

A species-rich genus with > 350 described species that occur in all zoogeographic regions except Antarctica. A molecular phylogeny of the genus was presented by Lin et al. (2018), placing Caladomyia as a junior synonym of Tanytarsus. The larvae are found in all types of freshwaters, with some marine, and at least one terrestrial species. The freshwater species usually construct long, soft tubes that are fixed to the bottom substrate (Epler et al. 2013).

Tanytarsus hastatus Sublette & Sasa, 1994: 56 [Guatemala]. Andersen et al. (2000) [Sinaloa State]; Vinogradova and Riss (2007) [“Yucatan Peninsula”]. European Bioinformatics Institute (2022). NT, NE. Brazil, Costa Rica, Ecuador, Guatemala, Mexico, Panama, Peru, USA, Venezuela.

Tanytarsus pistra (Sublette & Sasa, 1994: 54) (Caladomyia) [Guatemala]. Vinogradova and Riss (2007: 32) [“Yucatan Peninsula”]. NT, NE. Guatemala, Mexico, USA (Lathrop and Mulla 1995).

Genus Tribelos Townes, 1945

A genus with less than 10 named species distributed mainly in the Nearctic and Palaearctic regions. The genus is also recorded from the Neotropical region (Trivinho-Strixino et al. 2000). The larvae occur in littoral sediments of small to large water bodies (Epler et al. 2013).

Tribelos sp.: Contreras-Ramos and Andersen (1999: 4) [Campeche State].

Genus Xenochironomus Kieffer, 1921

A genus with ~ 20 species distributed in the Neotropical, Nearctic, Palaearctic, Oriental, and Australasian regions. The genus was revised by Fusari et al. (2013). The larvae of almost all species are obligate miners in freshwater sponges in standing and flowing waters (Epler et al. 2013).

Xenochironomus sp.: Contreras-Ramos and Andersen (1999: 4) [Campeche State]; Vinogradova and Riss (2007: 33) [“Yucatan Peninsula”].

Genus Xestochironomus Sublette & Wirth, 1972

A genus of more than ten described species that occur only in the Neotropical and Nearctic regions (Pinho and Souza 2013; Bello-González et al. 2016). Known larvae are miners in immersed wood in running waters (Epler et al. 2013).

Xestochironomus latilobus Borkent, 1984: 29 [Venezuela]. Andersen et al. (2000: 590) [Campeche State]; Contreras-Ramos et al. (2000: 26). NT. Costa Rica, Mexico, Venezuela.

Genus Zavreliella Kieffer, 1920

A genus with ~ 15 species; according to Fusari et al. (2017), 13 of these are known from tropical South America. The genus was revised by Reiss (1990). Larvae build transportable cases and move among submerged vegetation in standing water, but can also be found in sediments in flowing waters (Epler et al. 2013).

Zavreliella longiseta Reiss, 1990: 112 [Brazil]. Contreras-Ramos and Andersen (1999: 4, as Omisus sp.) [Campeche State]; Contreras-Ramos et al. (2000: 26, as Omisus sp.); Vinogradova and Riss (2007: 33) [“Yucatan Peninsula”]. NT. Brazil, Costa Rica, Mexico, Panama.

Remarks. The genus Omisus Townes, 1945 was recorded from Campeche State by Contreras-Ramos and Andersen (1999) and Contreras-Ramos et al. (2000). However, this record is incorrect. At closer examination the specimens belong to Zavreliella longiseta Reiss, 1990, a species that lacks dark spots in the wing and has a second, strong, curved spur on the hid tibia. The generic diagnosis given by Reiss (1990) should thus be amended accordingly.

Subfamily Diamesinae

Genus Diamesa Meigen, 1835

A genus of > 100 named species distributed in the Nearctic, Palaearctic, Afrotropical, and Oriental regions. Larvae of Diamesa are generally adapted to cool waters, inhabiting flowing water, springs, and to a lesser extent shallow still water and the hygropetric zone; they can be dominant in the kryon zone of glacier fed streams (Sæther and Andersen 2013a).

Diamesa mexicana Serra-Tosio, 1977: 100 [Mexico: Mexico State, Lake Nevado de Toluca]. Andersen et al. (2000: 589); Ashe and O’Connor (2009: 281). Endemic.

Diamesa reissi Serra-Tosio, 1977: 99 [Mexico: Mexico State, Lake Nevado de Toluca]. Andersen et al. (2000: 589); Ashe and O’Connor (2009: 283). Endemic.

Genus Pseudokiefferiella Zavřel, 1941

The only included species, Pseudokeifferiella parva (Edwards, 1932), is distributed in the Nearctic and Palaearctic regions. The larvae inhabit small streams and the hygropetric zone (Sæther and Andersen 2013a).

Pseudokiefferiella sp.: Bogan et al. (2014: 2726) [Sonora State].

Subfamily Orthocladiinae

Syn.: Prodiamesinae (see Lin et al. 2022).

Genus Allocladius Kieffer, 1913

A genus of 25 named species that occur in all zoogeographical regions, except Antarctica and Oceania. Andersen et al. (2010) reviewed the South American species; a revision of the genus was given by Ferrington and Sæther (2011). The larvae of Allocladius appear to be truly aquatic, as they have been found in ponds, rivers, and streams, including the shores of brackish water bodies and salt marshes, but some are probably able to survive in moist sandy substrata (Andersen et al. 2013b).

Allocladius nanseni (Kieffer, 1926: 82) (Camptocladius) [Canada]. Ferrington and Sæther (2011: 66) [Mexico State]; Ashe and O’Connor (2012a: 118). NE, PA. Widespread.

Genus Antillocladius Sæther, 1981

A genus of 30 named species that occur mostly in the Neotropical region, but are also found in the Nearctic, Palaearctic, and Oriental regions (Ashe and O’Connor 2012a; Andersen and Hagenlund 2017). The genus was reviewed by Mendes et al. (2004, 2011) and Mendes and Andersen (2008). Known larvae from South America appear to be terrestrial or semi-terrestrial as they have been collected in moss and lichens on stones and tree trunks; a North American species has been found in seeps near streams and impoundments (Mendes et al. 2004; Andersen et al. 2013b).

Antillocladius arcuatus Sæther, 1982: 474 [USA: South Carolina]. Mendes et al. (2004: 29) [Nuevo León State]; Mendes and Andersen (2008: 21); Ashe and O’Connor (2012a: 121). NT, NE. Brazil, Mexico, USA, Venezuela.

Antillocladius calakmulensis Mendes, Andersen & Sæther, 2004: 32 [Mexico: Campeche State, Calakmul Biosphere Reserve]. Mendes and Andersen (2008: 28); Ashe and O’Connor (2012a: 122); Admin (2022). Endemic.

Antillocladius herradurus Mendes, Andersen & Sæther, 2004: 39 [Mexico: Campeche State, Calakmul Biosphere Reserve]. Mendes and Andersen (2008: 33); Ashe and O’Connor (2012a: 122); Admin (2022). Endemic.

Antillocladius pluspilalus Sæther, 1982: 474 [USA: South Carolina]. Mendes et al. (2004: 48) [Campeche State]; Mendes and Andersen (2008: 36); Ashe and O’Connor (2012a: 122). NT, NE. Ecuador, Mexico, Nicaragua, USA.

Antillocladius zempoalensis Mendes, Andersen & Sæther, 2004: 57 [Mexico: Morelos State, Lagunas de Zempoala National Park]. Mendes and Andersen (2008: 41); Ashe and O’Connor (2012a: 124). Endemic.

Genus Bryophaenocladius Thienemann, 1934

A species-rich genus with ~ 120 named species that occur in all zoogeographic regions, except Antarctica and Oceania. Neotropical and Mexican species were reviewed by Wang et al. (2006). The larvae of most species are terrestrial or semi-terrestrial, but a few are aquatic (Andersen et al. 2013b).

Bryophaenocladius digitatus Sæther, 1973: 55 [USA: South Dakota]. Wang et al. (2006: 23) [Campeche State]; Ashe and O’Connor (2012a: 141). NE. Mexico, USA.

Bryophaenocladius humerosus Wang, Andersen & Sæther, 2006: 26 [Mexico: Morelos State, Lagunas de Zempoala National Park]. Ashe and O’Connor (2012a: 144); Admin (2022). Endemic.

Bryophaenocladius pichinensis Wang, Andersen & Sæther, 2006: 28 [Ecuador]. Wang et al. (2006: 28) [States of Nuevo León; Puebla]; Ashe and O’Connor (2012a: 150); Admin (2022). NT. Ecuador, Mexico.

Bryophaenocladius simplex Wang, Andersen & Sæther, 2006: 30 [Mexico: Nuevo León State, Allende, Río Ramos]. Wang et al. (2006: 30) [Nuevo León State, Santiago]; Ashe and O’Connor (2012a: 152); Admin (2022). Endemic.

Genus Cardiocladius Kieffer, 1912

A genus of 20 named species that occur in all zoogeographic regions except Antarctica and Oceania. The Neotropical species were reviewed by Andersen et al. (2016). The larvae live in fast-flowing waters and are often associated with the immature stages of blackflies (Simuliidae), on which they are reported to be predaceous (Andersen et al. 2013b).

Cardiocladius moreloensis Andersen, Hagenlund & Pinho, 2016: 277 [Mexico: Morelos State, Estación Ceamish]. Endemic.

Genus Clunio Haliday, 1855

A genus of 25 described species that occur in all zoogeographic regions except Antarctica. The larvae are marine and believed to be omnivorous, feeding on algae and dead or dying animals (Andersen et al. 2013b).

Clunio sp.: Sotelo-Casas et al. (2014: 17) [Nayarit State: Marieta Islands].

Genus Corynoneura Winnertz, 1846

A genus of ~ 100 named species that occur in all zoogeographic regions except Antarctica. A review of the Neotropical species was given by Wiedenbrug et al. (2012). Larvae occur in virtually all types of aquatic habitats, from standing waters to fast-flowing streams (Andersen et al. 2013b).

Corynoneura zempoala Wiedenbrug, Lamas & Trivinho-Strixino, 2012: 55. [Mexico: Morelos State, Parque Nacional Lagunas de Zempoala]. Endemic.

Genus Cricotopus Wulp, 1874

Syn.: Paratrichocladius Santos Abreu, 1918 (see Cranston and Krosh 2015)

A genus of ~ 270 named species that occur in all zoogeographic regions except Antarctica. Seven subgenera are recognized, namely Cricotopus s. str.; Isocladius Kieffer, 1909; Maurius Lehmann, 1981; Nostocladius Ashe & Murray, 1980; Oliveiriella Wiedenbrug & Fittkau, 1997; Paratrichocladius Santos Abreu, 1918; and Pseudocricotopus Nishida, 1987 (see Ashe and O’Connor 2012a; Andersen et al. 2013b; Cranston and Krosch 2015). Larvae inhabit all types of freshwaters including saline coastal waters. They are frequently associated with aquatic plants, including algae, and some mine living parts of aquatic macrophytes (Andersen et al. 2013b).

Cricotopus (Cricotopus) bicinctus (Meigen, 1818: 41) (Chironomus) [Austria]. Andersen et al. (2000: 589) [States of Mexico; Guerrero; Sinaloa]; Ashe and O’Connor (2012a: 209). NT, NE, PA, OR, OC. Widespread.

Cricotopus (Isocladius) sylvestris (Fabricius, 1794: 252) (Tipula) [Germany]. Andersen et al. (2000: 589) [States of Mexico; Guanajuato]; Ashe and O’Connor (2012a: 245). NT, NE, PA. Widespread.

Cricotopus (Cricotopus) triannulatus (Macquart, 1826: 202) (Chironomus) [France]. Andersen et al. (2000: 589) [Puebla State]; Ashe and O’Connor (2012a: 231); Alcocer et al. (2016: 411). NE, PA. Widespread.

Genus Diplosmittia Sæther, 1981

A genus of 10 named species distributed in the Neotropical and Nearctic regions. A review of the genus was provided by Pinho et al. (2009b). Wiedenbrug and Silva (2016) added a species from the Dominican Republic. The immatures are unknown.

Diplosmittia harrisoni Sæther, 1981: 30 [St. Lucia]. Pinho et al. (2009b: 177) [Campeche State]; Ashe and O’Connor (2012a: 262). NT. Costa Rica, Mexico, St. Lucia, St. Vincent, Venezuela.

Genus Gravatamberus Mendes & Andersen, 2008

A genus with five named species endemic to the Neotropical region. Larvae have been found in bromeliads (Mendes and Andersen 2008).

Gravatamberus curtus Mendes & Andersen, 2008: 45 [Mexico: Campeche State, Calakmul Biosphere Reserve]. Ashe & O’Connor (2012a: 293); Admin (2022). NT. Costa Rica, Mexico.

Remarks. Epler (2017) recorded Gravatamberus guatemaltecus Mendes & Andersen, 2008 from Zurquí de Moravia in Costa Rica and commented on the variation in G. curtus.

Genus Limnophyes Eaton, 1875

A genus of > 90 named species that occur in all zoogeographic regions except Oceania. Sæther (1990a, b) revised the Holarctic, Afrotropical, and Neotropical species of the genus. The larvae are eurytopic, including aquatic, semiterrestrial and terrestrial habitats (Andersen et al. 2013b).

Limnophyes sp.: Andersen et al. (2000: 591) [Puebla State].

Genus Lopescladius Oliveira, 1967

A genus with eight named species from the Neotropical and Nearctic regions. Two subgenera are recognized, namely Lopescladius s. str. and Cordiella Coffman & Roback, 1984 (see Ashe and O’Connor 2012a). South American species of Lopescladius (Cordiella) were described by Hagenlund et al. (2010). Larvae inhabit streams with sandy sediments (Trivinho-Strixino 2011).

Lopescladius (Lopescladius) verruculosus Sæther, 1983: 289 [Mexico: Michoacán State, Tocuman]. Andersen et al. (2000: 589); Ashe and O’Connor (2012a: 365). NE. Mexico, USA.

Genus Mesosmittia Brundin, 1956

A genus of 18 named species that occur in the Neotropical, Nearctic, Palaearctic, Afrotropical, and Oriental regions. The Neotropical and Mexican species were reviewed by Andersen and Mendes (2002b). The immatures are likely terrestrial (Andersen et al. 2013b).

Mesosmittia acutistylus Sæther, 1986: 43 [USA: New Mexico]. Andersen & Mendes (2002b: 143) [Campeche State]; Ashe and O’Connor (2012a: 369). NE. Mexico, USA.

Mesosmittia annae Andersen & Mendes, 2002b: 143 [Guatemala]. Andersen and Mendes (2002b: 143) [Campeche State]; Ashe and O’Connor (2012a: 369); Admin (2022). NT. Guatemala, Mexico.

Mesosmittia guanajensis Andersen & Mendes, 2002b: 147 [Mexico: Guanajuato State, Acámbaro]. Ashe and O’Connor (2012a: 370); Admin (2022). Endemic.

Mesosmittia lobiga Sæther, 1986: 45 [USA: New Mexico]. Andersen and Mendes (2002b: 150) [States of Guanajuato; Nuevo León]; Ashe and O’Connor (2012a: 370). NT, NE. Mexico, Puerto Rico, USA.

Mesosmittia patrihortae Sæther, 1986: 47 [USA: South Carolina]. Andersen and Mendes (2002b: 150) [States of Campeche; Nuevo León; Veracruz]; Ashe and O’Connor (2012a: 371). NT, NE, PA, AF. Widespread.

Remarks. Based on material collected in Zurquí, Costa Rica, Epler (2017) could not separate M. truncata from M. patrihortae Sæther, 1986, and considered M. truncata to be a junior synonym of M. patrihortae.

Mesosmittia prolixa Sæther, 1986: 48 [USA: Kansas]. Andersen and Mendes (2002b: 150) [States of Campeche; Nuevo León]; Ashe and O’Connor (2012a: 371). NE. Mexico, USA.

Mesosmittia tora Sæther, 1986: 50 [USA: South Dakota]. Andersen and Mendes (2002b: 150) [Nuevo León State]; Ashe and O’Connor (2012a: 371). NE. Mexico, USA.

Genus Metriocnemus Wulp, 1874

A genus of 75 named species that occur in all zoogeographic regions except Antarctica and Oceania. Three subgenera are recognized, namely Metriocnemus s. str.; Crymaleomyia Ashe & O’Connor, 2000; and Inermipupa Langton & Cobo, 1997 (see Ashe and O’Connor 2012a). A review of the genus was given by Sæther (1995). Larvae occur in mosses, phytotelmata, springs, ditches, streams and lakes and a few species are hygropetric (Andersen et al. 2013b).

Metriocnemus sp.: Andersen et al. (2000: 591) [Nuevo León State].

Genus Nanocladius Kieffer, 1913

A genus of 37 named species that occur in all zoogeographic regions except Antarctica. Two subgenera are recognized, namely Nanocladius s. str., and Plecopteracoluthus Steffan, 1965 (see Ashe and O’Connor 2012a). Neotropical species were treated by Wiedenbrug and Silva (2013). Larvae occur in streams, rivers, lakes, and ponds and some are symphoretic on immature Megaloptera and Ephemeroptera (Andersen et al. 2013b).

Nanocladius sp.: Contreras-Ramos and Andersen (1999: 4) [Campeche State]; Vinogradova and Riss (2007: 33) [“Yucatan Peninsula”].

Genus Onconeura Andersen & Sæther, 2005

A genus of eight named species that occur in the Neotropical and Nearctic regions. A review of the genus was given by Wiedenbrug et al. (2009), and a cladistic analysis of the genus was given by Donato et al. (2012). The larvae inhabit streams and rivers (Andersen et al. 2013b).

Onconeura semifimbriata (Sæther, 1981: 32) (Thienemanniella) [St. Vincent]. Andersen and Sæther (2005: 13) [Nuevo León State]; Wiedenbrug et al. (2009: 13); Ashe and O’Connor (2012a: 408). NT. Brazil, Costa Rica, Guatemala, Mexico, St. Vincent.

Genus Orthocladius Wulp, 1874

A genus of ~ 150 named species that occur in the Nearctic, Palaearctic, Afrotropical, and Oriental regions. Six subgenera are recognized, Orthocladius s. str., Eudactylocladius Thienemann, 1935; Euorthocladius Thienemann, 1935; Mesorthocladius Sæther, 2005; Pogonocladius Brundin, 1956; and Symposiocladius Cranston, 1982 (see Ashe and O’Connor 2012a). The genus is recorded from South America based on unnamed larvae from Argentina belonging to the subgenus Eudactylocladius (Wais 1987). The larvae inhabit all types of flowing waters, lakes, ponds, swamps, and moist earth; some species also mine submerged wood (Andersen et al. 2013b).

Orthocladius (Euorthocladius) sp.: Andersen et al. (2000: 591) [Mexico State].

Orthocladius (Orthocladius) sp.: Andersen et al. (2000: 591) [Mexico State].

Genus Paralimnophyes Brundin, 1956

A genus of five named species that occur in the Nearctic, Palaearctic, Oriental, and Australasian regions. The only species with described larvae inhabits eutrophic lowland pools and ditches (Andersen et al. 2013b).

Paralimnophyes sp.: Andersen et al. (2000: 591) [Puebla State]; Contreras-Ramos et al. (2000: 25) [Campeche State].

Genus Parametriocnemus Goetghbuer, 1932

A genus of 35 named species that occur in all zoogeographic regions except Antarctica and the Neotropical region. The genus is recorded from South America based on unnamed larvae from Brazil, Colombia, Peru, and Venezuela (Roback and Coffman 1983; Ospina-Torres et al. 1999; Trivinho-Strixino 2011). Larvae of Parametriocnemus are found in springs and in relatively fast flowing cold streams and rivers (Andersen et al. 2013b).

Parametriocnemus sp.: Contreras-Ramos and Andersen (1999: 4) [Campeche State]; Bogan et al. (2014: 2726) [Sonora State].

Genus Paratrichocladius Santos Abreu, 1918

See: Cricotopus Wulp, 1874.

Genus Prodiamesa Kieffer, 1906

A genus of six named species distributed in the Nearctic and Palaearctic regions. Larvae of Prodiamesa occur in springs, streams, rivers, ponds, and the littoral zone in lakes (Sæther and Andersen 2013b).

Prodiamesa sp.: Granados-Ramírez et al. (2017: 45) [Morelos State].

Genus Psectrocladius Kieffer, 1906

A genus with > 60 named species that occur in all zoogeographic regions, except Antartica, Australasia, Oceania, and the Neotropical region. Four subgenera are recognized, namely Psectrocladius s. str.; Allopsectrocladius Wülker, 1956; Mesopsectrocladius Laville, 1972; and Monopsectrocladius Wülker, 1956 (see Ashe and O’Connor 2012a). The only record from South America is an unnamed larval morphotype from the Peruvian Amazon belonging to subgenus Psectrocladius (Roback 1966). The larvae are eurytopic (Andersen et al. 2013b).

Psectrocladius sp.: Andersen et al. (2000: 591) [Puebla State]; Bogan et al. (2014: 2726) [Sonora State].

Genus Pseudosmittia Edwards, 1932

A genus of > 100 described species that occur in all zoogeographic regions, except Antarctica. Andersen et al. (2010) reviewed the Neotropical species, and a revision of the genus was given by Ferrington and Sæther (2011). Most larvae appear to be semiterrestrial to semiaquatic (Andersen et al. 2013b).

Pseudosmittia forcipata (Goetghebuer, 1921; 87) (Camptocladius) [Belgium]. Andersen et al. (2010: 39) [States of Campeche; Nuevo León]; Ferrington & Sæther (2011: 297); Ashe and O’Connor (2012a: 545). NT, NE, PA, OR. Widespread.

Pseudosmittia invirgata Andersen, Sæther & Mendes, 2010: 43 [Mexico: Campeche State, Calakmul Biosphere Reserve]. Ferrington and Sæther (2011: 288); Ashe and O’Connor (2012a: 547). Endemic.

Pseudosmittia joaquimvenancioi (Messias & Oliveira, 2000: 189) (Bryophaenocladius) [Brazil]. Wang et al. (2006: 19); Andersen et al. (2010: 45) [States of Campeche; Veracruz]; Ferrington and Sæther (2011: 184); Ashe and O’Connor (2012a: 547). NT. Brazil, Costa Rica, Mexico, Nicaragua, St. Lucia, St. Vincent, Venezuela.

Genus Rheocricotopus Brundin, 1956

A genus of ~ 75 described species that occur in all zoogeographic regions except Antarctica and Oceania. Two subgenera are recognized, namely Rheocricotopus s. str., and Psilocricotopus Sæther, 1986 (see Ashe and O’Connor 2012a). The first named species from the Neotropical region, Rheocricotopus (Psilocricotopus) sirventorum Andersen & Mendes, was recently described from Brazil by Andersen and Mendes (2012). Larvae are rheophilic, living on plants and stones in streams and rivers, and are rarely found in the littoral zone of lakes (Andersen et al. 2013b).

Rheocricotopus sp.: Andersen et al. (2000: 589) [Mexico State].

Genus Smittia Holmgren, 1869

A species-rich genus with > 80 named species that occur in all zoogeographic regions except Antarctica. Most larvae are terrestrial, occurring in damp soil, but at least one species is aquatic (Andersen et al. 2013b).

Smittia sp.: Andersen et al. (2000: 591) [Baja California Sur State]; Hamerlík et al. (2018: 217) [Yucatán State].

Genus Synorthocladius Thienemann, 1935

A genus of eight named species that occur in all zoogeographic regions except Antarctica. The larvae inhabit springs, small to large bodies of flowing water and small bodies or shallow parts of still water (Andersen et al. 2013b).

Synorthocladius semivirens (Keiffer, 1909: 48) (Dactylocladius) [Germany]. Andersen et al. (2000: 589) [Mexico State]; Ashe and O’Connor (2012a: 610). NE, PA, OR. Widespread.

Genus Thienemanniella Kieffer, 1911

A genus of ~ 55 named species that occur in all zoogeographic regions except Antarctica. The Neotropical species were reviewed by Wiedenbrug et al. (2013). The larvae occur in most lotic habitats, from fast-flowing streams to slow-flowing ditches and rivers (Andersen et al. 2013b).

Thienemanniella sp.: Contreras-Ramos and Andersen (1999: 4) [Campeche State]; Bogan et al. (2014: 2726) [Sonora State]; Granados Ramírez et al. (2017: 45) [States of Mexico; Morelos].

Subfamily Prodiamesinae

See Subfamily Orthocladiinae.

Subfamily Tanypodinae

Genus Ablabesmyia Johannsen, 1905

A genus of nearly 100 described species that occur in all zoogeographic regions, except Antarctica; it is currently the most speciose genus in Tanypodinae. Four subgenera, Ablabesmyia s. str., Asaya Roback, 1985, Karelia Roback, 1971, and Sartaia Roback, 1983 are recognized (see Ashe and O’Connor 2009). Most Neotropical species probably belong in Ablabesmyia s. str., but as pointed out by several authors, many South American species cannot be assigned to a subgenus with certainty, as there are inconsistencies in the establishment of these groups (see Neubern et al. 2013). Many of the recently described species are thus not assigned to a subgenus. The Neotropical species were reviewed by Neubern et al. (2013). The larvae occur in a wide variety of habitats, including small and large standing and flowing waters from cold temperate to warm tropical climate zones (Cranston and Epler 2013).

Ablabesmyia (Karelia) cinctipes (Johannsen, 1946: 271) (Pentaneura) [USA: Florida]. Andersen et al. (2000: 589) [States of Chiapas; Guerrero]; Vinogradova and Riss (2007: 33) [“Yucatan Peninsula”]; Ashe and O’Connor (2009: 121); Cranston and Epler (2013: 62). NT, NE. Bahamas, Belize, Guatemala, Mexico, St. Vincent, USA.

Genus Alotanypus Roback, 1971

A genus of 11 described species distributed in the Neotropical, Nearctic, Palaearctic, and Australasian regions. Larvae occur in both standing and flowing waters and appear to tolerate a broad range of conditions including very acid waters (Cranston and Epler 2013).

Alotanypus sp.: Andersen et al. (2000: 591) [Nuevo León State].

Genus Apsectrotanypus Fittkau, 1962

A genus of seven named species that occur in all zoogeographic regions except Antarctica and Oceania. In South America unnamed species are recorded from Argentina and Colombia (Donato et al. 2008b; Ruiz-Moreno et al. 2000; Spies and Reiss 1996). The larvae inhabit small, cool, flowing waters (Cranston and Epler 2013).

Apsectrotanypus sp.: Bogan et al. (2014: 2725) [Sonora State].

Genus Clinotanypus Kieffer, 1913

A genus of ~ 45 described species that occur in all zoogeographic regions, except Antarctica. Two subgenera are recognized, Clinotanypus s. str. and Aponteus Roback, 1971 (see Ashe and O’Connor 2009.) The Neotropical species were reviewed by Neubern et al. (2014). The larvae prefer soft sediments in shallow, warm water bodies including ponds, lakes and slow-flowing streams and rivers (Cranston and Epler 2013).

Clinotanypus sp.: Contreras-Ramos and Andersen (1999: 4) [Campeche State].

Genus Coelotanypus Kieffer, 1913

A genus of ~ 20 described species that occur in the Neotropical, Nearctic, Afrotropical, and Australasian regions. A key to the males of the Neotropical species was given by Paggi and Zilli (2018). The larvae inhabit benthic sediments of lakes, including artificial impoundments, slow flowing reaches of rivers and old riverbeds (Cranston and Epler 2013). The genus can be very abundant in Amazonian flood-plain lakes and in wetlands in southern Brazil (Fonseca Leal et al. 2004; Panatta et al. 2007).

Coelotanypus atus Roback, 1971: 37 [USA: Texas]. Andersen et al. (2000: 589) [Mexico, without specific locality]; Ashe and O’Connor (2009: 140). NT, NE. Mexico, Puerto Rico, USA.

Coelotanypus concinnus (Coquillett, 1895: 308) (Tanypus) [USA: Texas]. Andersen et al. (2000: 589) [Sonora State]; Ashe and O’Connor (2009: 141). NT, NE. Costa Rica, Mexico, Nicaragua, Puerto Rico, USA.

Coelotanypus naelis Roback, 1963: 170 [Surinam]. Andersen et al. (2000: 589) [Veracruz State]; Ashe and O’Connor (2009: 142). NT, NE. Mexico, Panama, Surinam, USA.

Coelotanypus olmecus Roback, 1965: 33 [Mexico: Veracruz State]. Andersen et al. (2000: 589); Ashe and O’Connor (2009: 142). NT. Mexico, Nicaragua.

Coelotanypus scapularis (Loew, 1866: 2) (Tanypus) [USA: Washington]. Andersen et al. (2000: 589) [Mexico, without specific locality]; Ashe and O’Connor (2009: 142). NT, NE. Canada, Mexico, Panama, USA.

Coelotanypus toltecus Roback, 1965: 32 [Mexico: Veracruz State]. Andersen et al. (2000: 589); Ashe and O’Connor (2009: 142). Endemic.

Coelotanypus tricolor (Loew, 1861: 309) (Tanypus) [USA: New York]. Andersen et al. (2000: 589) [Veracruz State]; Ashe and O’Connor (2009: 143). NT, NE. Costa Rica, Mexico, USA.

Genus Djalmabatista Fittkau, 1968

A genus of 15 described species that occur in all zoogeographic regions except Antarctica and Oceania. The larvae appear to prefer low alkalinity to weakly acid waters, and may be found in lakes, ponds, springs, large and small rivers, as well as in temperate to tropical lentic and lotic depositional habitats (Cranston and Epler 2013).

Djalmabatista pulchra (Johannsen, 1908: 273) (Protenthes) [USA: New York]. Andersen et al. (2000: 589) [States of Chiapas; Guerrero]; Ashe and O’Connor (2009: 155). NT, NE. Argentina ( Oca et al. 2020), Bahamas (Anderson et al. 2014), Brazil, Canada, Costa Rica, Guatemala, Mexico, Nicaragua, USA.

Genus Fittkauimyia Karunakaran, 1969

A genus of eight named species that occur in all zoogeographic regions except Antarctica and Oceania. The larvae inhabit rivers and the littoral zone of lakes, generally in tropical and subtropical regions (Cranston and Epler 2013).

Fittkauimyia sp.: Andersen et al. (2000: 591) [States of Campeche; Nuevo León]; Contreras-Ramos and Andersen (1999: 4); Contreras-Ramos et al. (2000: 25); Vinogradova and Riss (2007: 33) [“Yucatan Peninsula”]; Bogan et al. (2014: 2725) [Sonora State]; Hamerlík et al. (2018: 217) [States of Quintana Roo; Yucatán].

Genus Labrundinia Fittkau, 1962

A genus of ~ 40 named species distributed in the Neotropical, Nearctic, Palaearctic, and Oriental regions. The genus was revised by Silva et al. (2014). The larvae live in small, standing water bodies as well as in streams and rivers (Cranston and Epler 2013).

Labrundinia fosteri Roback, 1987: 2018 [Colombia]. Vinogradova and Riss (2007 33) [“Yucatan Peninsula”]; Ashe and O’Connor (2009: 164); Silva et al. (2014: 44). NT. Colombia, Mexico.

Labrundinia longipalpis (Goetghebuer, 1921: 66) (Tanypus) [Belgium].

Syn.: Labrundinia maculata Roback, 1971: 278 [USA: California] (Silva et al. 2011: 294).

Andersen et al. (2000: 589) [Coahuila State]; Ashe and O’Connor (2009: 165, 2012b: 127) [Michoacán State]; Silva et al. (2011: 295, 2014: 67). NT, NE, PA. Widespread.

Labrundinia pilosella (Loew, 1866: 5) (Tanypus) [USA: District Columbia]. Andersen et al. (2000: 589) [Puebla State]; Ashe and O’Connor (2009: 166); Silva et al. (2014: 127). NT, NE. Canada, Guatemala, Honduras, Mexico, Puerto Rico, Trinidad and Tobago, USA, Venezuela.

Genus Larsia Fittkau, 1962

A genus of ~ 30 named species that occur in all zoogeographic regions except Antarctica. Neubern and Silva (2011) described two new species from the Neotropical region and presented a checklist of the Larsia species of the world. In the Southern Hemisphere the larvae are associated with both lotic and lentic warm waters (Cranston and Epler 2013).

Larsia planensis (Johannsen, 1946: 284) (Pentaneura) [USA: Texas]. Andersen et al. (2000: 589) [Mexico City and States of Morelos; Oaxaca; Veracruz]; Ashe and O’Connor (2009: 169). NT, NE, OC. Canada, Guatemala, Hawaiian Islands, Mexico, USA.

Genus Natarsia Fittkau, 1962

A genus of six named species distributed in the Nearctic, Palaearctic, and Oriental regions. The larvae of the North American species live in small running waters, perhaps favoring cool water. European species inhabit streams, springs, and the littoral zone of montane or northern lakes and show hygropetric behavior in small, standing waters (Cranston and Epler 2013).

Natarsia sp.: Vinogradova and Riss (2007: 33) [“Yucatan Peninsula”].

Genus Nilotanypus Kieffer, 1923

A genus of 11 named species distributed in all zoogeographical regions except Antarctica and Oceania. Andersen and Pinho (2019) recently described two new species of Nilotanypus from Brazil. The larvae inhabit flowing waters, especially areas with sandy beds (Cranston and Epler 2013).

Nilotanypus sp.: Contreras-Ramos and Andersen (1999: 4) [Campeche]; Vinogradova and Riss (2007: 33) [“Yucatan Peninsula”].

Genus Paramerina Fittkau, 1962

See: Zavrelimyia Fittkau, 1962.

Genus Pentaneura Philippi, 1866

A genus of eight named species distributed in the Neotropical and Nearctic regions. Silva and Ferrington (2018) recently reviewed the Neotropical species. The larvae inhabit a variety of aquatic systems, from small streams and ponds to lakes and bays, occasionally the larvae live in shallow water flowing over bedrock covered with moss, algae, and detritus (Silva and Ferrington 2018).

Pentaneura inconspicua (Malloch, 1915: 371) (Tanypus) [USA: Illinois]. Andersen et al. (2000: 589) [Mexico City]; Ashe and O’Connor (2009: 195). NE. Canada, Mexico, USA.

Genus Procladius Skuse, 1889

The second most speciose genus of Tanypodinae, with ~ 70 named species that occur in all zoogeographical regions except Antarctica. Four subgenera are recognized, namely Procladius s. str., Holotanypus Roback, 1982, Laurotanypus Oliveira, Messias & Silva-Vasconcellos, 1992, and Psilotanypus Kieffer, 1906 (see Ashe and O’Connor 2009; Dantas and Hamada 2018). The larvae prefer muddy substrate of standing or slow-flowing water bodies, especially ponds and small lakes, but a few also inhabit the profundal zone of large, deep lakes (Cranston and Epler 2013).

Procladius (Psilotanypus) bellus (Loew, 1866: 4) (Tanypus) [USA: Washington]. Andersen et al. (2000: 589) [Mexico City]; Ashe and O’Connor (2009: 210); Bentley and Thomas (2022) [Puebla State]. NE, OR. Canada, China, Mexico, USA.

Procladius (Holotanypus) culiciformis (Linnaeus, 1767: 978) (Tipula) [Sweden]. Andersen et al. (2000: 589) [Mexico City]; Ashe and O’Connor (2009: 199). NE, PA. Widespread.

Remarks. The species was recorded from Campeche State by Mendoza-Arroyo and López-Toledo (2017: 27). This record is doubtful as the specimen was studied using a stereomicroscope with too low magnification to observe morphological details and no experts were involved in the identification of the specimen.

Genus Psectrotanypus Kieffer, 1909

A genus with seven named species that occur in the Nearctic, Palaearctic, Afrotropical, and Oriental regions. The genus was recorded from the Neotropical region by Fittkau and Reiss (1979), but without specifying a country. The larvae occur in ponds, bogs, small bodies of water and slow-flowing streams (Cranston and Epler 2013).

Psectrotanypus sp.: Alcocer et al. (2016: 411) [Puebla State].

Genus Tanypus Meigen, 1803

A genus of > 30 named species that occur in all zoogeographic regions except Antarctica and Oceania. Two subgenera are recognized, namely Tanypus s. str. and Apelopia Roback, 1971 (see Ashe and O’Connor 2009). The larvae live in sediments in standing and slowly flowing waters, especially in temperate to warm regions, where they can tolerate high salinity (Cranston and Epler 2013).

Tanypus (Tanypus) catemaco (Roback, 1964: 141) (Pelopia) [Mexico: Veracruz State]. Andersen et al. (2000: 589); Ashe and O’Connor (2009: 227). Endemic.

Tanypus (Apelopia) neopunctipennis Sublette, 1964b: 118 [USA: Illinois]. Andersen et al. (2000: 589) [States of Oaxaca; Veracruz]; Ashe and O’Connor (2009: 226). NT, NE. Bahamas, Cuba (Bello-González and Téllez-Martínez 2012), Mexico, USA.

Genus Thienemannimyia Fittkau, 1957

A genus of ~ 20 named species occurring in the Nearctic, Palaearctic, Afrotropical, and Oriental regions. Unnamed species were reported from Costa Rica by Watson and Heyn (1993). The larvae are found in both lotic and lentic waters (Cranston and Epler 2013).

Thienemannimyia sp.: Andersen et al. (2000: 591) [Nuevo León State].

Genus Zavrelimyia Fittkau, 1962

Syn.: Paramerina Fittkau, 1962.

Recently Silva and Ekrem (2016) formally placed the genus Paramerina Fittkau as a synonym of Zavrelimyia Fittkau. The genus now comprises ~ 50 named species that occur in all zoogeographic regions except Antarctica. Larvae of Zavrelimyia s. str. are, with few exceptions, more or less cold stenothermic and in temperate regions of the Holarctic primarily inhabitants of sandy or detritus rich sediments of springs and lentic habitats of stream sections close to springs. Larvae of Zavrelimyia (Paramerina) are eurythermic, living in a variety of standing waters of all sizes, but are also present in small lotic habitats including pools in rivers (Cranston and Epler 2013).

Zavrelimyia (Paramerina) smithae (Sublette, 1964b: 100) (Pentaneura (Pentaneura)) [USA: California]. Andersen et al. (2000: 589) [States of Oaxaca; Puebla]; Ashe and O’Connor (2009: 192). NE. Mexico, USA.

Generically unplaced valid Macropelopiini

roblesi Vargas, 1946: 80 (Macropelopia) [Mexico: Chiapas State, Mariscal]. Andersen et al. (2000: 589 as Macropelopia roblesi Vargas); Ashe and O’Connor (2009: 250, 362). Endemic.

Generically unplaced valid Tanypodinae

marmorata Johannsen, 1938: 219 (Pentaneura) [Puerto Rico]. Andersen et al. (2000: 589 as Pentaneura marmorata Johannsen) [States of Chiapas; Guerrero; Veracruz]; Ashe and O’Connor (2009: 252). NT. Mexico, Puerto Rico.

Subfamily Telmatogetoninae

Genus Telmatogeton Schiner, 1867

A genus of ~ 30 named species that occur in all zoogeographic regions. Except for a few freshwater species from Hawaii, Telmatogeton larvae are marine and live in the intertidal zone where they construct tubes within green algae such as Enteromorpha (Cranston and Ashe 2013).

Telmatogeton alaskensis Coquillett, 1900: 395 [USA: Alaska]. Andersen et al. (2000: 589) [Mexico, without specific locality]; Ashe and O’Connor (2009: 332). NE. Canada, Mexico, USA.

Telmatogeton latipenne Wirth, 1949: 172 [Mexico: Colima State, Revillagigedo Islands]. Andersen et al. (2000: 589); Ashe and O’Connor (2009: 333). Endemic.

Genus Thalassomya Schiner, 1856

A genus of 12 named species that occur in all zoogeographic regions except Antarctica. The larvae live in the intertidal marine zone, particularly in the warmer seas of the world (Cranston and Ashe 2013).

Thalassomya bureni Wirth, 1949: 167 [USA: Florida]. Andersen et al. (2000: 589) [Baja California Sur State]; Ashe and O’Connor (2009: 336). NT, NE. Mexico, USA. According to Wirth (1969) distributed “from Florida to Panama and the West Indies”.

Thalassomya longipes (Johnson, 1924: 86) (Galapagomyia) [Ecuador: Galapagos Islands]. Andersen et al. (2000: 589) [Nayarit State: Tres Marias Islands]; Ashe and O’Connor (2009: 337). NT. Ecuador, Mexico.

Thalassomya pilipes Edwards, 1928: 60 [American Samoa]. Andersen et al. (2000: 589) [Baja California State; Colima State: Revillagigedo Islands]; Ashe and O’Connor (2009: 338). NT, NE, OR, AU, OC. Widespread.

Species richness and taxonomic composition

A total of 110 species are listed for Mexico; 52 species in 25 genera belong to the subfamily Chironominae, 30 species in 13 genera to Orthocladiinae, 19 species in nine genera and two valid species that are not placed in a genus to Tanypodinae, five species in two genera to Telmatogetoninae, and two species in one genus to Diamesinae. In addition, there are records of 41 genera without identified species. Of these, 20 genera belong to Chironominae, 12 to Orthocladiinae, eight to Tanypodinae, and one genus to Diamesinae.

Distribution

The number of species recorded from the different states throughout Mexico is very uneven. More than ten species have only been recorded from six states. From Campeche a total of 29 species are recorded, most of them based on material collected during a project in Calakmul Biosphere Reserve (Contreras Ramos et al. 2000). From Veracruz 19 species have been recorded, from Nuevo León 15 species, from Puebla 13 species, from the State of Mexico 11 species and from Morelos ten species. From the remaining states only five or less species have been recorded. In most of the states in central and northern Mexico, as well as those on the Pacific coast, there are no or only a few records (Fig. 1).

Figure 1.

Biogeographic affinities and number of chironomid species recorded from each of the 32 Mexican states. Abbreviations: AC: Aguascalientes; BC: Baja California; BCS: Baja California Sur; CA: Chiapas; CC: Campeche; CDMX: Ciudad de México; CI: Coahuila; CM: Colima; CU: Chihuahua; DG: Durango; EM: Estado de México; GE: Guerrero; GJ: Guanajuato; HG: Hidalgo; JC: Jalisco; MA: Michoacán; MO: Morelos; NL: Nuevo León; NY: Nayarit; OX: Oaxaca; PA: Puebla; QR: Quintana Roo; QT: Querétaro; SI: Sinaloa; SL: San Luis Potosí; SO: Sonora; TB: Tabasco; TL: Tamaulipas; TX: Tlaxcala; VR: Veracruz; YC: Yucatán; ZC: Zacatecas.

The type localities for 34 Chironomidae species are in Mexico; of these, 27 species (25% of the total number of recorded species) are endemic. Twenty-nine species have a Neotropical distribution, 15 are Nearctic or Holarctic, while the remaining 39 species are distributed in both the Neotropical and Nearctic regions or are more widely distributed.

Discussion

In addition to being key to freshwater and riparian ecosystems (e.g., Porinchu and MacDonald 2003; Paetzold et al. 2005), chironomids have been widely used to recreate the environmental history of lakes and rivers (e.g., Plikk et al. 2019), generate typologies (e.g., Schöll and Haybach 2004; Nyman and Korhola 2005), propose biogeographical hypotheses (e.g., Brundin 1966; Krosch et al. 2011), ecotoxicological models (e.g., Beleza et al. 2019; Ferrari et al. 2019), biomonitoring (e.g., Gomes et al. 2018; Molineri et al. 2020) and for the evaluation of taxonomic and functional diversity (e.g., Jyväsjärvi et al. 2018). However, the Mexican Chironomidae fauna needs to be much better studied before it can be useful in such contexts.

In the previous checklist (Andersen et al. 2000), the number of species listed was 61; so, 49 species have been added during the last two decades. Of these, no less than 25 species belong to the subfamily Orthocladiinae, and the number of Orthocladiinae species has thus increased five times from the five species recorded in 2000. In Chironominae the number of species has increased from 29 species in 2000 to 52 species today; while in the subfamilies Tanypodinae, Telmatogetoninae, and Diamesinae no species have been added since 2000.

Comparing the number of Chironomidae species recorded in Mexico with the number in other neighboring, better studied areas, highlights the need for further studies in Mexico. Oliver et al. (1990) and Oliver and Dillon (1994) listed 206 generic and 1065 species names of Nearctic Chironomidae. More than 700 species of chironomids are listed from southeastern USA, including Alabama, Florida, Georgia, North and South Carolina, and Tennessee, which together comprise approximately 41% of the total area of Mexico (Caldwell et al. 1997). For the state of California, bordering Mexico and comprising ~ 22% of the area of Mexico, 245 species of chironomids have been recorded (Spies 1999). More than 400 species have been recorded from the state of Florida, which comprises less than 9% of the total area of Mexico (Epler 2019).

No comprehensive checklist for the Neotropical region has been published since Spies and Reiss (1996). However, in an updated checklist for Brazil (Pinho 2022) 658 species in 99 genera are listed. Mendes and Pinho (2016) recently published a checklist for Colombia listing only 30 species of Chironomidae in 16 genera in three subfamilies. In addition, 32 genera and two subfamilies have been recorded from Colombia based on larva, but without identified species.

The 110 species recovered in the present checklist is far from the 1000 species estimated by Andersen et al. (2000) to occur in Mexico and highlights the need for further studies. Most additional species to be found will undoubtedly belong to the subfamilies Chironominae, Orthocladiinae, and Tanypodinae. Chironominae is the most species rich subfamily of Chironomidae and is found in all biogeographical regions except Antarctica. Additional species will mainly be found in slow flowing streams and rivers, lakes and ponds in lowland habitats, but additional species will also be found in streams, rivers and lakes at higher altitudes. Mexican species of some genera, like e.g. Rheotanytarsus Thienemann & Bause, 1913 have been reviewed and new species described. However, there are several species-rich genera in which Mexican material has not or hardly been studied and in genera like Pseudochironomus Malloch, 1915, Tanytarsus Wulp, 1874, and Polypedilum Kieffer, 1912, many more species are likely to be added. The Orthocladiinae is also a very species-rich and widely distributed subfamily that tends to be particularly abundant in streams and rivers in mountainous areas. For some genera, like Antillocladius Sæther, 1981, Bryophaenocladius Thienemann, 1934 and Mesosmittia Brundin, 1956, Mexican material has been included in reviews of the genera, while other species-rich genera like Corynoneura Winnertz, 1846 and Cricotopus Wulp, 1874, are hardly studied at all. Most additional Tanypodinae species will probably be found in slow flowing streams and rivers, lakes and ponds in lowland habitats. So far only a few genera of Tanypodinae have been studied in detail in Mexico and for several species-rich genera like Ablabesmyia Johannsen, 1905 and Labrundinia Fittkau, 1962, there are only a few species recorded from Mexico so far.

Particularly in Orthocladiinae, several recently described genera like Colosmittia Andersen & Sæther, 1994, Litocladius Mendes, Andersen & Sæther, 2004, and Titimbera Andersen, Pinho & Mendes, 2015 might also occur in Mexico as they have all been taken in Costa Rica (Andersen et al. 2011a; Mendes et al. 2011; Andersen et al. 2015). There might well be several undescribed genera in the subfamily. Epler (2017) recently recorded no less than 16 undescribed genera of Orthocladiinae from Zurquí in Costa Rica.

Additional species will also likely be found in some of the less species-rich subfamilies. Today, ten extant subfamilies of Chironomidae are recognized. Six subfamilies occur in the Nearctic region, while in the Neotropical region no fewer than nine subfamilies have been encountered. At the subfamily level the Neotropical region is thus the most diverse biogeographical region. Only the monotypic subfamily Usambaromyiinae Andersen & Sæther has not been recorded. In the Neotropical region two of the other subfamilies, Chilonomyiinae Brundin and Aphroteninae Brundin, have only been found in southern Chile and Patagonia and it is unlikely that any species in these two subfamilies occur in Mexico.

However, two subfamilies so far not recorded from Mexico might occur in the country. The subfamily Buchonomyiinae Brundin & Sæther with three included species is found in the Neotropical, Palaearctic, and Oriental regions. It was recorded for the first time from the Neotropical region by Andersen and Sæther (1995) describing Buchonomyia brundini Andersen & Sæther, 1995 from a small, shallow, rather fast-flowing river in Costa Rica. The subfamily Podonominae Thienemann & Edwards has a mainly bipolar distribution with five genera and 15 species in North America and Canada, and five genera with altogether 85 species in the southern part of South America. Spies (1999) recorded two species of Boreochlus Edwards, 1938 and one species of Parochlus Enderlein, 1912 from California. Several species have recently also been described from Brazil, and two genera, Podonomus Philippi, 1866 and Parochlus, are listed from Colombia based on larvae (Mendes and Pinho 2016; Pinho 2022).

The subfamily Telmatogetoninae with two genera, Telmatogeton Schiner, 1867 and Thalassomya Schiner, 1856, is marine. Both genera with altogether five species are known from Mexico.

The subfamily Diamesinae has a mainly arctic or alpine distribution with 55 species in ten genera in the Nearctic region and 11 species in five genera in the Neotropical region. Two species of Diamesa Meigen, 1835 were described by Serra-Tosio (1977) from a high-altitude lake in the Mexico State. The genus, with 107 species, is known from the Nearctic, Palaearctic, Afrotropical, and Oriental regions. Spies (1999) listed six species of Diamesa from California and one species in each of the genera Pseudodiamesa Goethgebuer, 1939 and Sympotthastia Pagast, 1947. Based on larvae, Mendes and Pinho (2016) listed the genus Paraheptagyia Brundin, 1966 from Colombia. Paraheptagyia, with five species, is distributed in the southern part of the Neotropical region and two species occur in the Australasian region (Ashe and O´Connor 2009).

The uneven distribution of Chironomidae records throughout the states in Mexico clearly reflects the lack of Chironomidae studies. Some Nematocera groups are better studied than the Chironomidae in Mexico. Consideration of the general distribution patterns of these groups may suggest what can be expected for the chironomids. States like Oaxaca and Chiapas are among the richest when it comes to Culicidae, Simuliidae and Ceratopogonidae (Ibánez-Bernal and Coscarón 1996; Ibánez-Bernal et al. 1996). Bond et al. (2014) also demonstrated that the Pacific slope has a high diversity of aquatic insects. Climatic and topographic heterogeneity in southeastern Mexico leads to high environmental heterogeneity (Rodríguez et al. 2019). The area has a complex geology resulting in barriers such as the Isthmus of Tehuantepec that is responsible for increased diversity in several insect groups (Halffter and Morrone 2017). It is expected that future studies will show that the increase in the number of Chironomidae species will be particularly striking in Oaxaca and Chiapas.

Mexico is known to have a high proportion of endemic species. In well-studied groups like amphibians, reptiles, and mammals the proportion of endemic species is 60%, 51% and 31%, respectively (Hufnagel and Mics 2021). However, the number of records of chironomids from Mexico is clearly insufficient to appreciate patterns of endemism or clear biogeographic relationships.

To increase the number of species recorded from Mexico, taxonomic studies should be given priority. Even though rearing of larvae is important to associate the immatures with adults, chironomids are generally described based on adult males. To achieve an immediate increase in species numbers, further studies should thus focus on adults rather than on larvae and pupae. Fieldwork should be focused particularly on the states in central and northern Mexico, where the chironomid fauna is poorly known. The southeastern states along the Pacific coast should also be given special attention. Different habitats such as streams, rivers, lakes, and ponds should be visited, and collections should be made at different altitudes. Several chironomid species live in special habitats, like phytotelmata, and many species particularly among the Orthocladiinae, are semiterrestrial or terrestrial.

Acknowledgements

We are indebted to John Epler in Florida and to Martin Spies in Munich for information and assistance.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This research would not have been possible without funding from the Consejo Nacional de Ciencia y Tecnología de México (CONACyT) through a doctoral fellowship, the British Ecological Society (Small Research Grant SR21-1220), and the Tonolli Award from the International Limnological Society. Natura y Ecosistemas Mexicanos, A.C. also provided support for this research

Author contributions

Orestes C. Bello-González: compilation, updating and analysis of information, taxonomic review, writing of the manuscript, preparation of the figure. Trond Andersen: compilation, updating and analysis of information, taxonomic review, writing of the manuscript. Norman Mercado-Silva: writing of the manuscript.

Author ORCIDs

Orestes C. Bello-González https://orcid.org/0000-0002-7697-1618

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

Norman Mercado-Silva https://orcid.org/0000-0001-7764-8161

Data availability

All of the data that support the findings of this study are available in the main text.

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﻿Abstract

Key words

﻿Introduction

﻿Methods

﻿Citations for species names are arranged as follows

﻿Results

﻿Check list