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53. Diptera Chironomidae

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Freshwater Invertebrates
Insecta: Diptera, of the Malaysian Region
Chironomidae 711

Insecta: Diptera, Chironomidae

Peter S. Cranston
Department of Entomology, University of California, Davis, CA 95616, U.S.A.
(Email: [email protected])

INTRODUCTION
The Chironomidae, which are known popularly as non-biting midges, are a species-
rich family of flies, with over 5000 described species worldwide. The common name
of non-biting midges derives from the weak development of the mandibles in the
adult, in contrast to their close relatives, the biting midges or sandflies
(Ceratopogonidae) in which the female mouthparts are often designed for taking a
blood meal. The adult stage is short-lived, and only encountered in substantial
numbers when associated with emergence from shallow eutrophic waters when
they may cause nuisance, even inhalent allergies.
In the male, the plumose antenna with elongate apical flagellomere is characteristic,
and in both sexes the single branch of the wing vein M is diagnostic (Fig. 1). The
larvae are apneustic (lacking functional spiracles) with rare, non-regional exceptions
(Cranston et al. 1987). The larval prolegs are nearly always present, one pair on the
prothorax and another on the final abdominal segment. The head capsule is always
complete and prognathous (directed forwards), which differentiates from the family
Thaumaleidae. For 95% of aquatic species, separation from larval Ceratopogonidae,
can be made with the prolegs, which are paired and separate in most chironomids. A
few predominantly terrestrial larval Orthocladiinae that lack prolegs confuse the
situation, and it may be necessary to examine the pharyngeal apparatus which is
strong, with divergent arms in ceratopogonid larvae, but weak in Chironomidae.

GENERAL BIOLOGY
Although there are some non-aquatic larvae, about 95% of all species are aquatic, in
standing and flowing waters. Some larval chironomids are commonly called
bloodworms because they have the red blood pigment haemoglobin in their bodies
(e.g. Chironomus and Polypedilum), but not all chironomid larvae are red. The
presence of haemoglobin enables larvae to extract oxygen from extremely low oxygen
environments, such as those resulting from organic pollution. Larval chironomid
species track ecological conditions closely, and their distributions have long been
used to assess water quality. This topic is well reviewed by Johnson (1995). More
general biological information on the family, its systematics, ecology and impacts
upon humans are covered in Armitage et al. (1994).
712 Freshwater Invertebrates of the Malaysian Region

The Chironomidae are extremely important ecologically due to their abundance

and diversity and also the fact that the larvae, as well as the pupae and adults, are a
source of food for most macroinvertebrate and vertebrate predators. The larvae
exhibit an enormous variety of feeding types: predators (e.g. Tanypodinae), grazers,
detritivores, wood borers (e.g. Stenochironomus), leaf miners, filter feeders and
ectoparasites (e.g. Cardiocladius) (Dudgeon 1999). Xenochironomus burrows into
sponges upon which they feed, while yet other species are facultative or obligate
ectosymbionts (e.g. Epoicladius) and ectoparasites (some Nanocladius) or
commensals of or phoretic on other aquatic insects (Dudgeon 1999). Many chironomids
live in tubes which they construct from their own silk spun from salivary glands,
often incorporating diatoms, algae or fine sediment particles. Most Chironomid larvae
are micro-detritivores, some are algal/diatom grazers, and a few species, though
often abundant, filter particles from flowing water.

Life cycle
Adult Chironomidae are noticed mostly when the male flies form aerial swarms,
usually close to the breeding site. The females are less obvious, but they may be seen
when they fly into the male mating swarm, or when they return to water to lay their
eggs. The eggs are laid usually in a sticky string or ‘rope’ with many eggs enclosed in
a mucus or jelly. Oviposition sites range from all forms of running and standing
waters to containers such as tree holes, bamboo internodes, and other forms of
plant-contained waters (phytotelmata). There is no egg diapause, and hatching takes
place rapidly. Larvulae (newly hatched from the egg) are short-lived, dispersive by
drift, rarely encountered in survey, and difficult to identify.
All chironomids have four larval instars, whose increase in size between moults
follows Dyar’s law with remarkable precision. Most morphological features used in
identification are quite consistent between third and fourth instars, although ratios,
such as between antennal segment lengths, alter with age. Second instars are little
studied; where known they appear to possess intermediate features between larvulae
and later instars, and hence may be difficult to identify also. Since the major
contributions to biomass come from the 3rd, and more importantly the 4th instars,
undersampling of younger instars is unlikely to be a serious problem.
As a generalisation, temperate life histories vary from multivoltine to greater than
one year in duration. Life cycles in the tropics appear to be rapid, perhaps as little as
one week (e.g. Nolte 1995). Pupal exuvial sampling, involving intercept of the
exuviae left floating on the water surface after adult emergence, is a powerful tool to
examine tropical synchronicity and seasonality (e.g. Hardwick et al. 1995; Cranston
2000). Thus far there is little evidence for such effects which, together with observations
from individual rearings suggests that stochasticity determines larval densities and
emergence patterns in tropical lotic systems.
Insecta: Diptera, Chironomidae 713

REGIONAL TAXA
Of the currently recognized eleven subfamilies of Chironomidae (Cranston 1994),
only four are recorded with certainty from the region. Three of these, the Tanypodinae,
Orthocladiinae and Chironominae, are ubiquitous and virtually world-wide in
distribution. The immature stages of many species of the subfamilies Tanypodinae
and Chironominae tolerate elevated temperatures, and reduced oxygen levels in their
aquatic habitats and can be abundant in tropical waters. The Orthocladiinae contain
many taxa that are susceptible to elevated temperatures and reduced oxygen, and in
the tropics diversity may be reduced relative to temperate regions of the northern
hemisphere. Diversity of tropical Orthocladiinae is highest in well-oxygenated running
waters.
The Diamesinae, a predominantly cool-adapted subfamily with its greatest diversity
in the northern hemisphere, is represented in the region by a single species recorded
from high elevation (above 3000 m) on Mount Kinabalu in Sabah (Willassen 1988).
Based on their biology and distribution, it might be expected that other species exist
in high elevation areas in the region (Ashe 1990).
The unrecorded Telmatogetoninae are to be expected on marine shores, where
the larvae live amongst algae, especially on rocky shores where there is freshwater
seepage, but also in rock pools and algal mats on sandy shores. Telmatogeton japonicus
is virtually cosmopolitan and may have extended its distribution by maritime trade,
and might be expected in the region.
The distribution of the subfamily Podonominae is amphitropical, that is having
representatives in the cold to temperate regions of both hemispheres, being particularly
speciose in the Gondwanan land fragments (Brundin 1966). Current biogeography
indicates that podonomines are unlikely to be found in the region.
Of the remaining smaller subfamilies, perhaps only Buchonomyiinae might be
expected in the region. The sole genus, Buchonomyia Fittkau, includes a species
found in Myanmar. Chilenomyiinae is restricted to Chile, Usumbaromyiinae to central
Africa and Prodiamesinae to the temperate Holarctic region, thus making these taxa
unlikely to occur in the region on biogeographic grounds (Ashe 1990).
Within the dominant three subfamilies, Tanypodinae, Orthocladiinae and
Chironominae, most described taxa from the region can be allocated to genera that
occur outside the region. Many of these genera actually are quite widespread, even
cosmopolitan in their distribution. Examples of such speciose and ubiquitous genera
include Clinotanypus, Procladius, Ablabesmyia, Larsia, Nilotanypus, Paramerina
and Zavrelimyia amongst the Tanypodinae, Corynoneura, Cricotopus, Nanocladius,
Paraphaenocladius, Parametriocnemus and Thienemanniella amongst the
Orthocladiinae and among the Chironominae, Chironomus, Cryptochironomus,
Dicrotendipes, Microtendipes, Paratendipes, Polypedilum, Stictochironomus and
all genera of Tanytarsini. Genera of more restricted distribution include several
714 Freshwater Invertebrates of the Malaysian Region

Figure 1. A – adult male Chironominae; B – wing of Tanypodinae; C – wing of Diamesinae; D – wing

of Chironominae; E – distal tarsomeres of Diamesinae; F – distal tarsomeres of Tanypodinae; G – apex
of tibia of Orthocladiinae; H – apex of tibia of Chironominae; I – genitalia (right half, ventral) of
Chironominae; J – genitalia (right half, ventral) of Orthocladiinae. Abbreviations: ant, antenna; Cu, cubitus
(vein); fe, femur; gs, gonostylus; gx, gonocoxite; hyp, hypopygium; M, media (vein); R1–5, radius (vein);
ta4, tarsomere (4); ti, tibia.
Insecta: Diptera, Chironomidae 715

Chironomini described as new from Sumatran material by Kikuchi and Sasa (1990),
namely Asachironomus, Tobachironomus and Sumatendipes. At least the latter
genus is not endemic since it is a previously unrecognized synonym of the Afro-
Australian genus Conochironomus.
The lack of endemism in the list above may be somewhat misleading, because
undescribed taxa of generic rank have been mentioned, such as three genera of
Pentaneurini and five of doubtful placement within the Chironomini (Ashe 1990)
and some already described taxa do not closely match current generic concepts.
Material from Sungai Belalong in Brunei and some locations in Peninsular Malaysia
confirm the general pattern: many cosmopolitan genera, interspersed with few taxa
of uncertain taxonomic placement and more restricted distribution. Shangomyia
Sæther and Wang, was amongst these latter taxa but has recently been recognized
as widespread in the Oriental Region from India to south China (Cranston, 2003).
The discovery of the larval sites exemplifies a reason for some of the gaps in our
knowledge – the larvae mine immersed wood in rivers. Such ‘minor’ in-stream
habitats including wood, macrophyte leaves and stems, root masses and peat swamps
are very poorly studied, yet in the northern hemisphere and Australia they harbour a
diversity of interesting midges. Much aquatic biodiversity may be hidden in such places.
As noted by Ashe (1990), certain regional chironomid species, exemplified by
several species of Dicrotendipes reviewed by Epler (1988) are widely distributed,
perhaps even from India in the west to Australia, Micronesia and Japan in the east.
These observations are confirmed and extended in recent studies (mostly unpublished
observations of the author) to include species in the genera Larsia, Paramerina,
Pseudosmittia, Kiefferulus (including longilobus), and many species of
Polypedilum, including P. (Pentapedilum) convexum. Amongst the more extended
distributions, some species have been found to extend westwards as far as the
Afrotropical region. In other cases very similar taxa extending from Africa to Australia
have been shown to be subtly delimited within the range, e.g. Conochironomus
(Cranston and Hare 1995).

KEYS TO SUBFAMILIES AND GENERA

The keys that follow are organized in the first keys being to subfamily level, in each
life history stage, and following this are the keys to genus within each subfamily,
based on the larval stage. The larva is the stage most frequently encountered by
ecologists. Although for some faunas it is possible to provide keys to genus and
species for the pupal exuvial stage, in this region we have too little information to
provide such a key.
In using the keys it is important to remember that there are many chironomids,
and some will not fit the keys. This is because the author has not seen everything
from the region.
716 Freshwater Invertebrates of the Malaysian Region

Figure 2. A – larval Tanypodinae; B – head capsule, ventral, of Tanypodinae; C – head capsule, ventral,
of Chironominae; D – ligula and paraligulae of Tanypodinae; E – ventral surface of head capsule of
Tanypodinae; F – mentum of Chironominae; G – mentum of Orthocladiinae; H – mentum and prementum
of Diamesinae; I – antenna of Diamesinae; J – antenna of Orthocladiinae. Abbreviations: ant(1), antennal
segment (1); ap, anterior parapod; bl, blade; lig, ligula; mand, mandible; m, mentum; pc, procercus; plig,
paraligula; pp, posterior parapod; sm, submentum; S9, S10, ventral seta 9 and 10; Ssm, seta submenta;
vmp, ventomental plate, VP, ventral pit.
Insecta: Diptera, Chironomidae 717

Key to subfamilies: adults

1. Wing with cross vein M–Cu present (Fig. 1B,C) ................................................................. 2
- Wing with cross vein M–Cu absent (Fig. 1D) ....................................................................... 3

2. R2+3 branched into separate R2 and R3 (Fig. 1B), wing often microtrichiose. Tarsomere 4
simple (Fig. 1F) ................................................................................................... Tanypodinae
- R2+3 unbranched (Fig. 1C), wing bare. Tarsomere 4 cordiform (Fig. 1E) ............ Diamesinae
(One species Diamesa cranstoni recorded at high elevations from Mount Kinabalu)

3. Fore-leg with basitarsus longer than, or subequal to tibia (Fig. 1A). Spurs of mid- and hind-tibia
often comb-like, composed of fused spines (Fig. 1H). Male genitalia with gonostylus rather
inflexibly fused to gonocoxite (Fig. 1I) .............................................................. Chironominae
- Fore-leg with basitarsus shorter than tibia. Spurs of mid- and hin- tibia if comb-like, composed
of free spines (Fig. 1G). Male genitalia with gonostylus flexibly attached to gonocoxite (Fig. 1J)
.......................................................................................................................... Orthocladiinae

Key to subfamilies: larvae

1. Antenna retractile into head (Fig. 2B). Hypopharynx with distinctive toothed ligula (Fig.
2B,D), mentum usually weakly sclerotized (Fig. 2E) ......................................... Tanypodinae
- Antenna non-retractile (Fig. 2C). Mentum a strongly sclerotized plate (the main mouthpart)
(Fig. 2F,G), with hypopharynx lacking strong ligula ............................................................. 2

2. Mentum associated with variably developed, but always broad and usually striated ventromental
plates (Fig. 2F) ................................................................................................... Chironominae
- Mentum without, or at most with relatively small, non-striate ventromental plates (Fig. 2G)
............................................................................................................................................... 3

3. Central ligula and lateral paraligulae of hypopharynx resembling three brushes (Fig. 2H). Third
antennal segment annulate (Fig. 2I) ........................................................................ Diamesinae
- Prementum never developed as three “brushes”. No annulate antennal segments (Fig. 2J) ....
.......................................................................................................................... Orthocladiinae

Key to subfamilies: pupae

1. Anal lobe fringed with setae, but without any distinctive marginal macrosetae (Fig. 3A,B).
Postero-lateral corner of segment VIII usually with spur or comb (Fig. 3A,C,D). Thoracic horn
often complex, bi- multi-branched to plumose (Fig. 3A) ................................... Chironominae
- Anal lobe with or without fringe, when fringed with distinctive marginal macrosetae (Fig. 3F).
Postero-lateral corner of segment VIII without spur or comb. Thoracic horn simple (Fig. 3E) or
absent ..................................................................................................................................... 2

2. Thoracic horn with internal horn sac and apical plastron plate (Fig. 3E). .......... Tanypodinae
- Thoracic horn without visible horn sac and lacking plastron plate ........................................ 3

3. Dorsomedian thorax with 3 setae, with the most posterior in a supralar position (Fig. 3H) ..
............................................................................................................................... Diamesinae
- Dorsomedial thorax with four setae, none in supralar position (Fig. 3I) .......... Orthocladiinae
718 Freshwater Invertebrates of the Malaysian Region

Figure 3. A – pupa of Chironominae; B – tergite VIII and anal lobe of Chironominae; C, D – spur of
segment VIII of Chironominae; E – thoracic horn of Tanypodinae; F, G – tergite VIII and anal lobe of
Orthocladiinae; H – thorax of Diamesinae; I – thorax of Orthocladiinae. Abbreviations: dc, dorsocentral
setae of thorax; ms, macrosetae of anal lobe; th, thoracic horn.
Insecta: Diptera, Chironomidae 719

SUBFAMILY TANYPODINAE
Key to genera: larvae
1. Body segments bearing longitudinal lateral dense setal fringe. Dorsomentum distinct, toothed
(Fig. 4A) ................................................................................................................................. 2
- Body segments lacking setal fringe. Dorsomentum indistinct, not toothed (Fig. 4B) .............
................................................................................................................. tribe Pentaneurini...6
2. Dorsomental teeth not aligned on plate (Fig. 4C). Mandible strongly hooked, with large basal
tooth (Fig. 4D) .................................................................................................... Clinotanypus
- Dorsomental teeth at margin of plate (Fig. 4E). Mandible straight to curved, never hooked ... 3

3. Pecten hypopharyngis absent. Mandible squat, with very expanded base and short apical tooth
(Fig. 4F) ...................................................................................................................... Tanypus
- Pecten hypopharyngis present. Mandible longer and more slender ......................................... 4

4. Mandible with multiple spine-like teeth on mesal surface, without distinct basal tooth (Fig. 4G)
............................................................................................................................ Fittkauimyia
- Mandible without subapical spine-like teeth ............................................................................. 5
5. Antennal blade extending well beyond apex of flagellum (Fig. 4H). Bases of anterior parapods
connected by region of fine hooklets (Fig. 4J) ................................................... Djalmabatista
- Antennal blade not extending beyond flagellum (Fig. 4I). Anterior parapods not connected by
spinose area ............................................................................................................ Procladius
6. Basal segment of maxillary palp divided into two or more sections (Fig. 5A) .......................... 7
- Base of maxillary palp undivided (Fig. 5B) .............................................................................. 8
7. One or more claws of posterior leg darker than remainder (Fig. 5C). Basal palp segment may be
divided into 2–5 sections; if only 2 sections, then membranous part anterior to basal 1/3. Submentum
with SSm nearly aligned with VP and S10, with S9 anteromedial to alignment (Fig. 5D) .............
............................................................................................................................ Ablabesmyia
- All hind proleg claws yellow. Membranous division of maxillary palp about 1/3 from base.
Submentum with seta S9 linearly aligned with SSm and S10, with VP lying more posterolateral
(Fig. 5E) ................................................................................................................ Paramerina
8. Apex of antenna with segment 3 as long as segment 2, with Lauterborn organs as long as
segment 3, darkened and partly fused at base to segment 2, giving “tuning-fork” arrangement
(Fig. 5F) ................................................................................................................................. 9
- Apex of antenna with segment 3 shorter than segment 2, with Lauterborn organs short, pale, and
not fused basally (Fig. 5G) ..................................................................................................... 11
9. Teeth of ligula forming straight row (Fig. 5H) .................................................. Telmatopelopia
- Teeth of ligula forming concave row (Fig. 5I,J) ..................................................................... 10
10. Middle section of ligula “waisted” (constricted). Middle tooth of ligula small relative to inner
tooth (Fig. 5I) ...................................................................................................... Monopelopia
- Middle section of ligula not constricted; middle tooth of ligula subequal to inner tooth (Fig. 5J)
........................................................................................................................... Krenopelopia
720 Freshwater Invertebrates of the Malaysian Region

Figure 4. A – mentum of Procladius sp.; B – mentum and submental setae of Telmatopelopia; C –

mentum of Clinotanypus; D – mandible of Clinotanypus; E – mentum of Fittkauimyia; F – mandible of
Tanypus; G – mandible of Fittkauimyia; H – antenna of Djalmabatista, I – antenna of Procladius; J –
anterior parapods of Djalmabatista.
Insecta: Diptera, Chironomidae 721

11. Inner teeth of ligula apically curved outwards (Fig. 5K). Submental setae S9 and S10 lying
adjacent, just anterior to VP (Fig. 5L) ........................................................ Thiennemannimyia
- Inner teeth of mentum directed anteriorly or slightly inwards. Submental setae S9 and S10
never in this formation ......................................................................................................... 12

12. Ligula with middle tooth larger than inner teeth (Fig. 6A). Submentum with setae (S9, S10 and
SSm) and VP linearly arranged with SSm distantly posterior to anterior group (Fig. 6B) ......
.............................................................................................................................. Nilotanypus
- Ligula with middle tooth subequal or shorter than inner (Fig. 6C). Submental setae and VP
variously arranged, never linear; SSm less distant from S9, S10 and SSm ........................... 13

13. Submentum with setae S9 anterior to VP, S10 antero-lateral to VP (Fig. 6D). Gula and submentum
smooth, unsculptured ................................................................................................... Larsia
- Submentum with seta S10 generally postero-lateral to VP .................................................. 14

14. VP lying medial to remaining setae. Gula and submentum with two or three transverse creases
(Fig. 6E) ................................................................................................ Pentaneurini Kelian C
- VP lying amongst remaining setae. Gula and submentum +/- smooth ................................ 15

15. SSm level with VP - setae on transverse +/- linear alignment (Fig. 6F) ...................................
.............................................................................................................. Pentaneurini Kelian B
- SSm posterior to VP - setae in triangular arrangement (Fig. 6G) .......... Pentaneurini Kelian A

SUBFAMILY TANYPODINAE
Late instar larvae are predatory. Many genera have species that are tolerant of
pollution, particularly high organic loadings (e.g. Procladius, amongst the most
pollution tolerant of chironomids, and also Clinotanypus, Tanypus, Fittkauimyia
and Paramerina). Some genera such as Procladius, prefer lentic conditions and
some (e.g. Tanypus) are common in warm, shallow pools. Nilotanypus is often
very abundant and widespread in sandy-bedded streams and rivers of Southeast
Asia and Australia. Larsia is a very common genus, especially in warm streams
and rivers running over sand. Paramerina is amongst the most common lotic
tanypodines in Southeast Asia. Although three species have been described from
the region, they cannot yet be related to material available to me. However, one
species, P. ignobilis is distinctive in the adult stage by the dark patterned wing, and
in its biology: it is found in treeholes in Southeast Asia, and in northern Queensland
rainforest.

SUBFAMILY DIAMESINAE
The subfamily Diamesinae is represented in the region by only one species, Diamesa
cranstoni, collected at high elevation on Mount Kinabalu, Sabah (Willassen 1988).
722 Freshwater Invertebrates of the Malaysian Region

Figure 5. A – palp of Ablabesmyia (hilli); B – palp of Thienemannimyia; C – posterior parapod claws

of Ablabesmyia; D – submentum of Ablabesmyia (hilli); D – submentum of Paramerina; E – apex of
antenna of Monopelopia; F – apex of antenna of Thienemannimyia; G – ligula of Telmatopelopia; H
– ligula of Monopelopia; I – ligula of Krenopelopia; J – ligula of Thienemannimyia; K – submentum
of Thienemannimyia.
Insecta: Diptera, Chironomidae 723

Figure 6. A – ligula of Nilotanypus; B – submentum of Nilotanypus; C – ligula of Larsia; D –

submentum of Larsia; E – submentum of Kelian A; F – submentum of Kelian B; G – submentum of
Kelian C.
724 Freshwater Invertebrates of the Malaysian Region

Figure 7. A, B – generalized larvae; C – mentum of Rheocricotopus; D – mentum of Parakiefferiella;

E – mentum of Nanocladius; F – head of Corynoneura; G – antenna of Corynoneura; H – antenna of
Thienemanniella. Abbreviations: ap, anterior parapods, m, mentum, pc, procercus, pp, posterior
parapods, vmp, ventromental plate.
Insecta: Diptera, Chironomidae 725

SUBFAMILY ORTHOCLADIINAE
Key to genera: larvae
1. Procercus absent, although procercal setae may indicate site. Parapods may be small, absent or
fused; claws usually present (Fig. 7B) ....................................... terrestrial and marine species
- Procercus present, apically bearing procercal setae. Parapods may be basally fused but are
always distinct, with claws (Fig. 7A) ................................................................................... 2
2. Ventromental plate well developed, extending laterally beyond lateral margin of outermost
tooth of mentum (Fig. 7C–E) ................................................................................................ 3
- Ventromental plate weaker to indistinguishable, never extending beyond lateral margin of
outermost mental tooth (may extend postero-lateral to base of outermost tooth)(Fig. 8A,B)
............................................................................................................................................... 5

3. Setae beneath ventromental plate (Fig. 7C). SI seta bifid ............................... Rheocricotopus
- No setae beneath ventromental plate. SI seta never bifid. ..................................................... 4

4. Ventromental plate extending just lateral of outermost mental tooth (Fig. 7D) .......................
........................................................................................................................ Parakiefferiella
- Ventromental plate extending more strongly lateral and posterior to base of outermost mental
tooth (Fig. 7E) ...................................................................................................... Nanocladius
5. Antenna about half head length or longer (Fig. 7F) ................................................................ 6
- Antenna shorter than half head length ................................................................................... 7
6. Antenna 4- segmented, longer than head (Fig. 7F, G) ......................................... Corynoneura
- Antenna 5-segmented (Fig. 7H), shorter than head ...................................... Thienemanniella
7. Ventromental plate with complex sinuous sclerotisation extending basal or basolateral to outer
mental tooth, posterior to base of outermost tooth (Fig. 8A) ................... Parametriocnemus
- Ventromental plates simple, linear, with single postero-lateral curvature, at most (Fig. 8B) 8
8. Antenna 4- segmented (Fig. 8C) ............................................................................................ 9
- Antenna 5- segmented .......................................................................................................... 10
9. No long procercal seta. Subapical seta of procercus strong (Fig. 8D) ................ Eukiefferiella
- One procercal seta extremely long, up to half body length. Subapical seta of procercus weak
(Fig. 8E) .............................................................................................................. Krenosmittia
10. Procercus short. Premandibles and ungula strongly sclerotized and robust (Fig. 8F) .............
.......................................................................................................................... Cardiocladius
- Procercus normally developed. Premandible and ungula not so robustly developed .......... 11
11. Abdominal segments posterolaterally with complex setal tuft (Fig. 8G). Mandible often with
outer margin distinctly crenulate, sometimes with inner margin spinose (Fig. 8H).... Cricotopus
- Abdominal setae simple. Mandibular margins smooth ........................................................ 12
12. S1 seta of labrum simple (Fig. 8I) ................................................................... Paracricotopus
- S1 seta of labrum bifid (Fig. 8J) .................................................................. Paratrichocladius
726 Freshwater Invertebrates of the Malaysian Region

Figure 8. A – mentum of Parametriocnemus; B – mentum of Eukiefferiella; C – antenna of Eukiefferiella;

D – procercus of Eukiefferiella; E – posterior segments of Krenosmittia; F – dorsal labrum of
Cardiocladius; G – lateral seta of Cricotopus; H – mandible of Cricotopus; I – labrum of Paracricotopus;
J – labrum of Paratrichocladius. Abbreviations: pmd, premandible, SI, anterior labral seta S.
Insecta: Diptera, Chironomidae 727

SUBFAMILY ORTHOCLADIINAE
Larvae of Eukiefferiella Krenosmittia, Paracricotopus, Parakiefferiella,
Parametriocnemus, Paratrichocladius and Rheocricotopus are all collector-gatherers
feeding on fine particulate organic matter. Cricotopus larvae graze on diatoms and
algae, Nanocladius may be ectoparasitic but more conventionally are associated with
filamentous algae. Corynoneura larvae are collector-gatherers or micro-filterers, while
Thienemanniella larvae are collector-gatherers that probably micro-filter.
Cardiocladius are associated with simuliid larvae in fast-flowing water where they
may be predatory on simuliids. Cricotopus species range from pristine-streams only,
to habitats with high levels of nutrients and algal production. Species of Eukiefferiella
appear to tolerate elevated water temperatures, and organic loadings, while species of
the genus Parametriocnemus in northern Australia are very tolerant of acidic
mine drainage (Hardwick et al. 1995). Thienemanniella larvae are very small and so
easily overlooked, but they are amongst the most common orthoclads in tropical streams.

SUBFAMILY CHIRONOMINAE
Key to genera: larvae
1. Bases of SI setae fused and SII on long pedestal (Fig. 9A). Antenna always with 5 segments
with Lauterborn organs distinct, sometimes on pedestals, never alternate .......... Tanytarsini .. 2
- Bases of SI rarely fused, SII rarely on pedestal (Fig. 9B). Antenna may be six segmented and
Lauterborn organs alternate, though usually opposite, sometimes indistinct ............................. 7
2. Ventromental plates separated medially by width of three median mental teeth (Fig. 9C) ....... 3
- Ventromental plates separated medially by no more than the width of the median mental tooth
(Fig. 9D) ................................................................................................................................. 4
3. Lauterborn organs alternate on segment 2; antennal pedestal with apical, non-palmate process
(Fig. 9E) ............................................................................................................. Stempellinella
- Lauterborn organs opposite, apical on segment 2; antennal pedestal with medial palmate process
(Fig. 9F) ................................................................................................................. Stempellina
4. Premandible with more than two teeth (Fig. 9A) .................................................................... 5
- Premandible bifid (Fig. 9G) ..................................................................................................... 6

5. Second antennal segment cylindrical, longer than third. Smaller Lauterborn organs arise from
long pedicels (Fig. 9H) ............................................................................................ Tanytarsus
- Second antennal segment distinctly wedge-shaped, subequal in length to third segment. Large
Lauterborn organs arise from short pedicels (Fig. 9I) .................................... Cladotanytarsus

6. Lauterborn organs on pedestals. Mandible with pronounced outer hump (Fig. 9J) ........ Sublettea
- Lauterborn organs sessile. No mandibular hump ............................................. Rheotanytarsus
7. Ventromental plates without striae, with few hooks (Figs. 10A,B) .......................................... 8
- Ventromental plates distinct, striate (Fig. 9C,D) ...................................................................... 9
728 Freshwater Invertebrates of the Malaysian Region

Figure 9. A – labrum of Cladotanytarsus; B – labrum of Cladopelma; C – mentum and ventromental

plates of Stempellina; D – mentum and ventromental plates of Rheotanytarsus; E – antenna of
Stempellinella; F – antenna of Stempellina; G – labrum of Rheotanytarsus; H – antenna of Tanytarsus;
I – antenna of Cladotanytarsus; J – mandible of Sublettea.
Insecta: Diptera, Chironomidae 729

8. Head dorso-ventrally flattened. Mentum more or less concave (Fig. 10B).... Stenochironomus
- Head not dorso-ventrally flattened. Mentum with strongly projecting median tooth/teeth (Fig.
10A) ..................................................................................................................... Shangomyia
9. Mentum with deep median V-shaped cleft, with slight indications of serrations on antero-median
margin (Fig. 10C) .................................................................................................. Fissimentum
- Mentum never cleft as above ................................................................................................ 10
10. SI seta plumose, SII not blade-like. Labral lamellae present. Pecten epipharyngis usually a wide
plate, rarely reduced to three sometimes serrate lobes (Fig. 10D) ......................................... 11
- SI and SII setae simple, frequently blade-like. Labral lamellae absent. Pecten epipharyngis usually
reduced to single scale, rarely toothed (Fig. 10E) ............................ Harnischia complex ... 21
11. All teeth of mentum and premandible pale. Head bean-shaped in lateral profile (Fig. 10F) .......
.............................................................................................................................. Nilothauma
- At least lateral mental teeth and apical mandibular teeth dark. Profile of head not bean shaped
............................................................................................................................................. 12
12. Antenna 5- segmented with Lauterborn organs either opposite on 2nd segment or absent (Fig.
10G) ...................................................................................................................................... 13
- Antenna 6- segmented with Lauterborn organs alternate, either on apices of 2nd and 3rd
segments or alternate on 2nd segment (Fig. 10H) ................................................................. 16

13. Mandible with basal striations (Fig. 11A). Generally with two pairs of ventral tubules and one
pair of lateral tubules although some reduction may occur .................................... Chironomus
- Mandible lacking basal striations, although exceptionally if present, without any abdominal tubules
............................................................................................................................................. 14
14. Ventromental plates narrow, only about 2x as wide as deep (Fig. 11B) .............. Dicrotendipes
- Ventromental plates broad, at least 2.5x as wide as deep (Fig. 11C) ..................................... 15
15. Premandible with at least 5 teeth. Pit on anterior of frons (Fig. 11D) ..................... Kiefferulus
- Premandible with 2–3 teeth. Frons without pit ...................................................... Polypedilum
16. Ventromental plates subtriangular, almost in medial contact (Fig.11E). Antenna subequal in length
to head (Fig. 11F) .................................................................................................... Zavreliella
- Ventromental plates more semilunar, more widely separated medially. Antenna shorter than head
............................................................................................................................................. 17
17. Median part of mentum pale (Fig. 11H,I) .............................................................................. 18
- Median part of mentum dark (Fig. 12A,B) ............................................................................ 19
18. Median mentum with four small pale teeth (Fig. 11H). Basal Lauterborn organ in middle of
second segment (Fig. 11G) .................................................................................. Paratendipes
- Median mentum with two or three pale teeth (Fig. 11I). Basal Lauterborn organ on apex of
second segment (Fig. 10G) ............................................................................... Microtendipes
19. Uneven number of subequal mental teeth (Fig. 12A) .................................... Stictochironomus
- Even number of disparate-sized mental teeth (Fig. 12B) ..................................................... 20
730 Freshwater Invertebrates of the Malaysian Region

Figure 10. A – mentum of Shangomyia; B – mentum of Stenochironomus; C – mentum of Fissimentum;

D – labrum of Chironomus; E – labrum of Paracladopelma; F – head of Nilothauma; G – antenna of
Stenochironomus; H – antenna of Paratendipes.
Insecta: Diptera, Chironomidae 731

20. Lauterborn organs on alternate apices of segments 2 and 3 (Fig. 12C) ........ Conochironomus
- Basal Lauterborn organ in mid-third antennal segment (Fig. 12D) .............................. Skusella
21. Mentum concave with pale median tooth (Fig. 12E) ............................................................. 22
- Mentum conventionally convex (Fig. 12F) ........................................................................... 23
22. Antenna 5-segmented. Blade on subapical segment 2 (Fig. 12G) .............. Cryptochironomus
- Antenna 6-segmented. Blade subapical on segment 3 (Fig. 12H) .............................................
.......................................................................................... Demicryptochironomus (Irmakia)
23. Mentum with outermost group of 3 teeth forming group distinctively set-off from general slope
(Fig. 12F,K) ........................................................................................................................... 24
- Mentum generally with teeth decreasing evenly in height from median to outermost (Fig. 12L)
............................................................................................................................................. 25
24. Mentum with single median tooth; ventromental plate with crenulate or spinose anterior margin
(Fig. 12F) ..................................................................................................... Microchironomus
- Mentum with paired median teeth; ventromental plate with smooth anterior margin (Fig. 12K)
............................................................................................................................. Cladopelma

25. Antenna with some hyaline sections, probably 6-segmented, but with indistinct segmentation;
style arising from middle of second segment (Fig. 12I) .......... Paracladopelma and Saetheria
- Antenna with 5 distinct segments, lacking hyaline sections; bifid style arising at apex of first
segment (Fig. 12J) ......................................................................................... Parachironomus

SUBFAMILY CHIRONOMINAE
Tribe Tanytarsini
Cladotanytarsus larvae are collector-gatherers that can attain high abundances in
sediment-rich rivers. Tanytarsus larvae are collector-gatherers. Larvae of
Rheotanytarsus are considered to be filterers, producing silk mesh between the
protruding arms of their tubes. Rheotanytarsus is a world-wide genus of many species,
with a taxonomy based especially on the genitalia of the male adult, and on the very
diagnostic pupal exuvial pattern. Larvae are much more difficult to distinguish.
Cladotanytarsus is a world-wide genus with many species, but with a poorly-defined
taxonomy. Tanytarsus is a world-wide genus with many species, with a taxonomy
based especially on the genitalia of the male adult, and on the very diagnostic pupal
exuvial pattern. Larvae of all Tanytarsini are difficult to distinguish, and in every
region studied, there is a mis-match between the number of larval types recognized
and the total number of species present.

Tribe Chironomini
Chironomus is one of the largest, and most difficult genera of chironomids. Adult
morphology is rather homogeneous, and we have not made much progress in
separation of larvae. Larvae are collector-gatherers, and are often found in organically
732 Freshwater Invertebrates of the Malaysian Region

Figure 11. A – mandible of Chironomus; B – mentum and ventromental plates of Dicrotendipes; C –

mentum and ventromental plates of Kiefferulus; D – labrum of Kiefferulus; E – mentum and ventromental
plates of Zavreliella; F – antenna of Zavreliella; G – antenna of Paratendipes; H – mentum of
Paratendipes; I – mentum of Microtendipes.
Insecta: Diptera, Chironomidae 733

Figure 12. A – mentum of Stictochironomus; B – mentum of Conochironomus; C – antenna of

Conochironomus; D – antenna of Skusella; E – mentum of Cryptochironomus; F – mentum of
Microchironomus; G – antenna of Cryptochironomus; H – antenna of Demicryptochironomus; I –
antenna of Paracladopelma/Saetheria; J – antenna of Parachironomus; K – mentum of Cladopelma;
L – mentum of Paracladopelma.
734 Freshwater Invertebrates of the Malaysian Region

enriched aquatic environments, both lotic and lentic. Cladopelma is a world-wide

genus with one species recorded from the region: Cladopelma viridula. The larvae
are quite common in a variety of water bodies, including rice fields. The larvae are
considered to be collector-gatherers. Conochironomus larvae are found in fine
sediments in slow-flowing rivers, pools, and lakes. They appear to be collector-
gatherers. Cryptochironomus is a world-wide and abundant genus.
Cryptochironomus larvae are found in fine to coarse sediments in slow-flowing
rivers, pools, and lakes. They appear to be collector-gatherers, but may also be
predatory. Dicrotendipes is one of the few genera that have been adequately studied
in the region (Epler 1988). Three species are recognized: all have wide distributions
in the region, extending to Australia in the south and southern Japan in the north.
Dicrotendipes larvae are fairly common in running waters, and will tolerate quite
high nutrient levels. Dicrotendipes larvae are collector-gatherers as are those of
Fissimentum and Microtendipes.
The larvae of Kiefferulus are collector-gatherers found in fine sediments,
especially in ponds, pools (including those in drying rivers) and in rice paddies.
Kiefferulus longilobus is found in saline inland and coastal waters where the flies
may become a nuisance (Cranston, Webb and Martin 1990). Microchironomus is a
small, but widely distributed genus. The species M. tener is one of the most widely
distributed species of chironomid, extending from Europe to Australia, and to Africa
in the south. The larva is found in standing waters, including some with quite high
levels of salinity. Microchironomus larvae are considered to be collector-gatherers.
Nilothauma larvae are found only in high quality waters.
The genus Parachironomus is widespread, but nowhere is it ever abundant.
The larvae of Parachironomus, and also those of Paracladopelma, are collector-
gatherers and may be predatory in some species. Paratendipes is a widespread
genus, but the taxonomy is very poorly known. Although the larvae are quite easy to
recognize (according to the characters in the key), the pupa and adult tend to merge
with other genera that also have a six-segmented antenna and alternate Lauterborn
organs in the larval stage.
Polypedilum is the most species-rich genus of Chironominae, and is very common
and diverse in tropical waters, including those of Borneo, Sumatra, Java and Bali.
Polypedilum larvae are collectors gatherers, found in all kinds of water bodies,
including phytotelmata. P. convexum is found in the latter habitat, including within
Nepethes pitcher plants. Polypedilum watsoni is abundant in the organically-polluted
reaches of the Kelian River, Kalimantan.
Insecta: Diptera, Chironomidae 735

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