Acta zoologica cracoviensia,46(suppl.– Fossil Insects): 381-392, Kraków, 15 Oct., 2003
Main ecological events in aquatic insects history
Nina D. SINITSHENKOVA
Received: 20 March, 2002
Accepted for publication: 30 April, 2002
SINITSHENKOVA N. D. 2003. Main ecological events in aquatic insects history. Acta zoo-
logica cracoviensia,46(suppl.– Fossil Insects): 381-392.
Abstract. In the Carboniferous no one undoubtedly aquatic insect remain is found. May-
flies and stoneflies were the oldest aquatic insects found in the Early Permian. Permian
was the time when the aquatic insects became diverse and probably colonized different
types of waters. Only one mayfly is found in the Early Triassic. The Middle and Late Tri-
assic aquatic insect assemblages are characterized by combination of Paleozoic and
Mesozoic elements. Diverse and widespread lacustrine insects are typical for the Jurassic.
Recent families of aquatic insects appeared in the Early Cretaceous. The Late Cretaceous
assemblages are considerably impoverished. The Cenozoic is the time of recolonization
of lacustrine habitats.
Key words: aquatic insects, history, Carboniferous, Permian, Triassic, Jurassic, Creta-
ceous, Cenozoic, palaeoentomology.
Nina D. SINITSHENKOVA. Palaeontological Institute, Russian Academy of Sciences, ul.
Profsoyuznaya 123, 117997 Moscow GSP-7, Russia.
Aquatic insects play most important role in modern fresh water ecosystems. Different, phyloge-
netically unrelated taxa with different types of development (both Holo- and Hemimetabola) are as-
sociated with aquatic environments. They differ in feeding habits as well, including detritivores,
herbivores and predators.
It is necessary to stress also the differences in the breathing mechanisms. Immature stages of the
mayflies, stoneflies, megalopterans, caddisflies, as well as some dipterans and beetles breathe with
dissolved oxygen, while adults and larvae of most beetles and bugs and immatures of some dipter-
ans breathe with atmosphere air and hence do not depend on the oxygen concentration in water. It is
plausible to assume that they had different patterns of colonization of aquatic environments because
of very different required adaptations.
The statement that the insects are secondarily aquatic animals, which had evolved from terres-
trial ancestors seems to be out of any serious doubt now. It is demonstrated that the wingless insects
existed in terrestrial habitats as early as in the Devonian. The oldest finds of winged insects come
from the Early Carboniferous (Namurian A) while the earliest undoubtedly aquatic insects appear
only in the Permian.
For recognizing aquatic insects among the fossils WOOTTON (1988) has proposed three criteria.
The main evidence of the aquatic mode of life he considered the presence of special adaptations for
aquatic life, such as swimming legs, gills and fringed caudal filaments. Less reliable criteria are
based on phylogenetic affinities of fossils with present-day aquatic taxa.
WOOTTON considers relation of a given fossil to a group originated from an aquatic ancestor to
be more reliable indication than when it belongs to a group with an uncertain way of life. The
WOOTTON’s criteria should be supplemented by the taphonomic ones, first of all by repeated fossil
finds of immature stages for which the probability of burial if they were terrestrial is low. It is the
most unreliable criterion. For instance, the aquatic mode of life has been suggested for some Per-
mian grylloblattids when basing on their frequent finds. Recent investigation of their body structure
demonstrated characteristic terrestrial features, so their aquatic life has been rejected completely
(STOROZHENKO 1998). It is necessary to take in consideration also the presence of ichnofossils in
the case when their classification to a definite group is well argued.
Acknowledgements.Thework is supported by RFBR 01-05-64741 and
II. CARBONIFEROUS TO PERMIAN
No Carboniferous insects demonstrating any obvious adaptations to aquatic mode of life are
known. As far as the taxonomic criteria are concerned, the Carboniferous mayfly-like Syntonopte-
rida and dragonflies of the suborder Meganeurina could be proposed. For the Carboniferous nymph
Bojophlebia (KUKALOVA-PECK 1985) an aquatic habit is doubtful. KLUGE (1996) considers it to be
a terrestrial insect related to Tysanura rather than a mayfly-like nymph.
Adult Meganeurina are abundant in the Carboniferous but complete absence of their nymphs in
the fossil record made PRITYKINA (1980) to suggest terrestrial habits for the Paleozoic odonatans.
An aquatic or semiaquatic way of life seems to be probable for Dasyleptidae – a peculiar extinct
group of the order Machilida occasionally common both in the Carboniferous and Permian. No
modern bristletails are aquatic, and dasyleptids demonstrate no special morphological features for
living inhabiting in water; however, their common presence in the fossil state is unique among the
apterygotans. The ichnofossils from the Late Carboniferous estuary deposits of Kansas were sup-
posedly attributed to dasyleptids (MÁGNANO et al. 1997). If this attribution is correct, Dasyleptidae
could prove to be the only Carboniferous insect family whose aquatic mode of life is quite probable.
When taking in consideration that it belongs to the order that is exclusively terrestrial now, there is
no continuity between the Carboniferous and post-Paleozoic aquatic insect faunas.
In any case the insects did not play any essential role in the Carboniferous fresh water ecosys-
tems. This fact fits well the general opinion about pronounced differences between those ecosys-
tems and the modern ones; however, the available data on insects do not allow to specify those
differences in a more precise way.
The first unequivocally aquatic immatures of winged insects enter fossil record in the Permian.
They include the mayfly and stonefly nymphs. However, their finds still remain rare in comparison
with the fossil adults of the same groups. The Early Permian mayfly nymphs (Fig. 1) are found in
Oklahoma, USA, in the Czech Republic, Central Europe (HUBBARD &KUKALOVA-PECK 1980),
and in the Urals in the famous locality Tshekarda (TSHERNOVA 1965). They represent two extinct
families, Protereismatidae in America and Central Europe, and Misthodotidae in Russia. Their
laminated abdominal gills (or tergaliae) and fringed caudal filaments indicate an aquatic mode of
Several stonefly nymphs of the family Tshekardoperlidae (Fig. 2) are collected in Tshekarda to-
gether with the misthodotid mayflies (SINITSHENKOVA 1987). The tshekardoperlid adults still re-
main undiscovered. Numerous adult stoneflies found in the same locality are assigned to other
families - Palaeonemouridae and Perlopseidae (SINITSHENKOVA 1987). Tshekardoperlid nymphs
were probably carnivorous. They had no gills, and their aquatic mode of life can be postulated only
on the base of their general similarity with modern perloid stoneflies. However, in Tshekarda an-
N. D. SINITSHENKOVA
other stonefly nymph is found, which shows quite obvious adaptations for inhabiting fast running
water (the rhithral zone). This is Barathronympha victima (Fig. 3) having the streamlined body, the
cerci moved apart widely, the flattened and widened femora and numerous short hairs on the tarsi
lacking the claws (SINITSHENKOVA 1987). Those characters are unique among the stoneflies, so that
the nymph resembles superficially a heptageniid mayfly rather than a stonefly.
One more remarkable nymph of uncertain systematic position is described from Tshekarda. Syl-
vonympha tshekardensis possesses the thoracic gills similar to the coxal gills of stoneflies (Fig. 4)
but cannot be assigned to this order and seems to represent an extinct lineage (NOVOKSHONOV &
Main ecological events in aquatic insects history 383
Figs 1-2. The Early Permian aquatic insects. 1 – mayfly Kukalova americana DEMOULIN, Protereismatidae, Oklahoma,
USA (after KUKALOVA-PECK 1968); 2 – stonefly Sylvoperlodes zhiltzovae SINITSH., Tshekardoperlidae, Urals, Russia
(after SINITSHENKOVA 1987).
Figs 3-4. The Early Permian aquatic insects of Urals, Russia. 3 – stonefly Barathronympha victima SINITSH., Nemouromor-
pha inc. sed., (after SINITSHENKOVA 1987); 4 – Sylvoperlodes tshekardensis NOVOKSH.etPAN’KOV, Perlidea, (after
NOVOKSHONOV &PAN’KOV 1999).
In Tshekarda insects occur in deltaic deposits of a large river. The mayflies and tshekardoperlid
stoneflies could inhabit this river, while Barathronympha dwelled most probably in a small rapid
The caddisflies appear in the Early Permian as well but they are represented exclusively by fos-
sils of adults, and the biology of their larvae is a matter of speculations. If they had inhabited some
protected microhabitats in running waters like many modern annulipalpian larvae do, they had little
if any chance to be preserved as fossils. In younger deposits, including the Cenozoic, the lotic cad-
disflies are represented by adults and occasionally by the integripalpian larval cases but the annuli-
palpian larval fossils are totally absent.
Dasyleptids still survived in the Early Permian; they are rather common in some areas (e.g., in
Kansas, USA) but never occur together with other aquatic insects. The most striking feature of the
Early Permian aquatic insects is that they never occur in lacustrine sediments, unlike the younger
deposits. The Early Permian finds seem to be restricted to deltaic and estuarine paleoenvironments.
The diversity of insects is usually very low, except the Tshekarda assemblage.
Further enrichment of the aquatic entomofauna is documented in the Late Permian (Fig. 5). The
mayflies are represented exclusively by winged stages. Some of them are related to the Early Per-
mian Misthodotidae (KINZELBACH &LUTZ 1984), while others resemble modern ephemeroids and
may belong to this lineage (MARTYNOV 1931). In the both cases nymphal aquatic mode of life is
quite probable, supposedly in large rivers.
Stoneflies occur in the Late Permian more often than in the Early Permian, but the finds of adults
still dominate. However, in the lacustrine deposits of West Siberia and Kazakhstan the stonefly
nymph remains are sometimes quite common. This fact permitted to propose their invasion to
standing waters. It is clear that in the Late Permian the stoneflies were diverse, widely distributed
and inhabited, probably, different types of water bodies (SINITSHENKOVA 1987).
Megalopterans are the only holometabolan insect order represented in the Late Permian by both
adults and larvae (Fig. 6) with characteristic abdominal gills (SHAROV 1953; PONOMARENKO 1976,
Fig. 5. The Permian sites with aquatic stages of insects. – Early Permian; – Late Permian.
N. D. SINITSHENKOVA
The schizophoroid beetles appear in the Late Permian and at once in many localities of Europe,
Asia, Australia, South America and South Africa. The presence on their elytrae of a special closing
device (the so-called “schiza”) resembling that of living aquatic beetles permitted PONOMARENKO
(1969) to suggest aquatic life for them. If so, schizophoroids are the oldest pterygotans with aquatic
adults. Their larvae are unknown.
Like in the Early Permian, the caddisflies are not rare but represented only by adults. The young-
est Dasyleptidae are known from the Late Permian of West Siberia. They are restricted to the only
locality Kaltan where stonefly nymphs occur as well in fine-grained sediments (RASNITSYN 2000).
Thus, in the Permian the aquatic insects become diverse and probably colonized gradually dif-
ferent types of waters, both running and standing. The occurrence of the Carboniferous and Early
Permian finds of aquatic stages and ichnofossils in estuarine and deltaic deposits seems to confirm
the statement of WOOTTON (1972) that the insects inhabited at first the running waters. However, in
my opinion this assumption seems rather disputable. Life in running waters suggests considerable
level of specialization including diverse morphological adaptations (e.g., to fixing on the substrate)
as well as specialized behavior with upstream flights of adults to compensate the drift of immature
stages. Hence true lotic insects should be rather advanced in comparison with early water coloniz-
In my view, periodically flooded habitats are the most probable biotopes of ancestral aquatic in-
sects; this scenario was postulated earlier by PONOMARENKO (1996). Many terrestrial insects de-
void of evident special adaptations can survive temporarily submersion. For them the colonization
of aquatic environments should be rather simple and gradual. This way of colonization may be
called a passive one. If such a passive colonization took place, then it is not surprising that the oldest
Main ecological events in aquatic insects history 385
Fig. 6. The Late Permian dobsonfly Permosialis sp., Orenburg region, Russia (after SHAROV 1953).
aquatic insects occur just in deltaic and estuary paleoenvironments where temporary flooding must
have been a common event. Appearance of morphologically highly specialized rheophilous
nymphs in the Early Permian indicates an early expansion from near-shore habitats to the riverbed.
Possibly, an important factor of such specialization was the protection of long-developing imma-
tures from the terrestrial and air predators. It is noteworthy that flying adults of the most ancient
aquatic groups – e.g., the stoneflies, megalopterans and especially mayflies – became short living.
The WOOTTON’s idea that the earliest aquatic insects were carnivorous (WOOTTON 1972) gives
rise to doubt. In particular, the Carboniferous dasyleptids were probably detritivorous. In the Per-
mian the groups with different feeding types are known, including detritivores and possibly grazing
algophages (mayflies, some stoneflies, possibly schizophoroid beetles) as well as predators (some
In the Early Triassic few insect finds are known, and only one mayfly adult has been recently
found in Siberia. Hence all our knowledge refers to the Middle and Late Triassic (Fig. 7). The dasy-
leptids are absent in the Triassic but all insect orders which appeared in the Permian not only sur-
vived but also even become more numerous and wider distributed.
Several orders are adding to the aquatic insect list in the Triassic, including the aquatic bugs and
dipterans as well as the most ancient undoubtedly aquatic odonatan nymphs in the Upper Triassic of
Australia (ROZEFELDS 1985).
Among the Triassic mayflies there are both the Permian survivors (Misthodotidae) and typically
Mesozoic groups (Mesoneta BRAUER,REDTENBACHER,GANGLBAUER, 1889; Mesobaetis BRAUER,
REDTENBACHER,GANGLBAUER, 1889) (SINITSHENKOVA 2000). The largest and richest Triassic
mayfly assemblage is discovered in the Grés-à-Voltzia deposits of the Vosges, France (MAR-
CHAL-PAPIER 1998). At present seven species are preliminary recognized as the nymphs, all but one
Fig. 7. The Triassic sites with aquatic stages of insects.
N. D. SINITSHENKOVA
previously undescribed. The most interesting are two species with unusually strong cuticle suggest-
ing that they possibly could survive in wet substrates at the shore zone. The oldest burrowing may-
fly nymphs are found as well.
Triassic stoneflies are widespread but represented mainly by adults. The nymphs are rather un-
common; some Late Triassic nymphs are assigned to several genera common in the Jurassic
The aquatic bugs are represented by rare Triassocoridae and supposed Naucoridae in the Middle
and Late Triassic of Ukraine, Central Asia and Australia (POPOV 1980; POPOV, pers. comm.). The
oldest numerically rich aquatic bug assemblage is discovered in the Uppermost Triassic of North
America (the Caw Branch Formation) (OLSEN et al. 1978; FRASER et al. 1996).
The undescribed megalopteran larvae with well-developed abdominal gills are found in France
and Ukraine (KALUGINA 1980). The schizophoroid beetles are common and diverse in the Triassic
over the World, and the oldest hydradephagan beetles are appearing at that time (PONOMARENKO
1969; ARNOLDI et al. 1977). The caddisflies are represented exclusively by adults.
Appearance of aquatic dipterans was likely the most important Triassic novelty in the aquatic in-
sects fauna if the importance of the order in all younger aquatic ecosystems is taken into account.
Dipteran larvae and pupae are found in Vosges Mts but not studied yet (MARCHAL-PAPIER 1998).
In spite of the relative scarcity of data, it is possible to say with certainty that the Triassic was the
time of considerable radiation of aquatic insects, both lotic and lentic. This radiation suggests that
the hydrological and hydrochemical regime of standing water bodies became more stable in com-
parison with the Paleozoic. In particular, in the Late Triassic the oldest lotic assemblages closely re-
sembling some widespread Jurassic types (e.g., with abundant aquatic bugs and with the
mesoleuctrid stonefly nymphs) are discovered.
The Jurassic insects are studied much better but mainly in North Asia. In Europe the Jurassic in-
sects are found principally in allochthonous oryctocenosis in marine deposits. In the North America
and Gondwanaland only few localities are known. All aquatic orders known in the Triassic are pres-
ent in the Jurassic, and are considerably more diverse.
The mayflies, stoneflies and odonatans belong mostly to extinct families (Fig. 8-10). The diver-
sity of the former two orders is higher in more temperate interior regions of Asia while diversity of
the odonatans culminates likely in warmer central Asia and Europe (TSHERNOVA 1967, 1969, 1977;
The aquatic bugs are diverse and widely distributed. The aquatic beetles are common and their
diversity is also rather high; the role of hydradephagan beetles is increasing while the schizophor-
oids decline gradually. The hydradephagan larvae are not rare and include both benthic and actively
swimming nectic forms (PONOMARENKO 1995).
In the Early Jurassic the recent mecopteran family Nannochoristidae appears in palaeontologi-
cal record for the first time. This is the only living aquatic group within the order. The nannochoris-
tid wings are not uncommon but the larval fossils are extremely rare (NOVOKSHONOV 1997). Their
rarity suggests that they probably dwelled in small streams like their modern relatives do. Nanno-
choristid larvae were active predators inhabiting running waters.
The Jurassic caddisflies are represented mainly by the primitive and probably unnatural annuli-
palpian family Necrotauliidae but there are other families as well including some primitive integri-
palpians. In the Upper Jurassic of Siberia, Mongolia and North America the caddisfly larval cases
appear in paleontological record for the first time (SUKATSHEVA 1985). Since that time they became
a basic component of lacustrine benthos. It is interesting to note that the case-building larvae are
algal-feeders and detritivores, with very few predators.
Main ecological events in aquatic insects history 387
Figs 8-9. Stoneflies from Early or Middle Jurassic of Siberia, Russia (after SINITSHENKOVA 1987). 8 – Platyperla platypoda
BRAUER,REDTENBACHER,GANGLBAUER 1889, Platyperlidae; 9 – Mesoleuctra tibialis SINITSH. Mesoleuctridae.
Fig. 10. Mayfly Epeoromimus kazlauskasi TSHERNOVA, Epeoromimidae, Early or Middle Jurassic of Siberia, Russia (after
N. D. SINITSHENKOVA
Since the Early Jurassic the nematoceran dipterans (in particular Chaoboridae and Chironomi-
dae) became extremely common in the lacustrine deposits (KALUGINA 1980).
Thus, in the Jurassic the taxonomic and ecological diversity of aquatic insects, especially in
lakes, are considerably higher in comparison with the Triassic. Some Jurassic freshwater ecosys-
tems were likely basically similar to recent ones, e.g. the ecosystems of oligotrophic cold-water
montane lakes and streams. On the other hand, we failed to find any modern analogs for some other
Jurassic ecosystems, e.g. for the small oxbow lakes of North Asia (SINICHENKOVA &ZHERIKHIN
The Cretaceous aquatic insects are well represented in the paleontological record both in the
northern continents and in Gondwanaland. The Early Cretaceous aquatic insect assemblages are di-
verse and taxonomically rich. They often include the families and even genera common in the Juras-
sic, and that is why the age of some deposits is disputable. On the contrary, the Late Cretaceous
assemblages are extremely impoverished and remarkably uniform. It should be stressed that since
the Cretaceous the insects occur in the fossil resins (ambers) that provides a valuable source of in-
formation about the running water fauna.
Among the mayflies the family Hexagenitidae is often most abundant , both in the north hemi-
sphere (Asia) (TSHERNOVA 1980) and in the south hemisphere (Brazil) (MCCAFFERTY 1990). Dif-
ferent families dominate some Asiatic assemblages and the Koonwarra fauna in Australia (JELL &
DUNCAN 1986). The burrowing mayfly nymphs are abundant in some sites. No lentic mayflies are
known from the Late Cretaceous while the lotic fauna was rather diverse as indicated by resin inclu-
sions from Asia and North America.
In comparison with the Jurassic the role of heterophlebioid and isophlebioid odonatans (the so-
called Mesozoic anisozygopterans which, however, had a little in common with the living Epio-
phlebiidae) decreases considerably, and the radiation of anisopteran dragonflies of the suborder Li-
bellulina is observed. The dragonflies are represented mostly by wings. Their nymphs may occur in mass
in lacustrine deposits but are always much less diverse than adults, which suggests that the majority
of species developed outside large lakes (PRITYKINA 1980; JELL &DUNCAN 1986; BECHLY 1998).
The species diversity of the Early Cretaceous dragonflies obviously decreased from the warm re-
gions in favor of the more temperate ones, just like it was in the Jurassic. The true zygopteran dam-
selflies are extremely rare. In the Late Cretaceous the odonatan fossils are represented nearly
exclusively by adults.
The stoneflies become uncommon and only rarely occur in considerable numbers in Cretaceous
deposits. However, their total diversity seems to increase in the Early Cretaceous because the main
Jurassic families still occur and some Recent ones enter the fossil record for the first time. All eco-
logical types of nymphs which existed in the Jurassic are known from the Early Cretaceous. Till
now no stonefly remain from the Late Cretaceous is found (SINITSHENKOVA 1987).
Aquatic bugs are common, diverse and widespread in the Early Cretaceous, especially the co-
rixids and notonectids. Unlike many other aquatic insects, bugs occur in Late Cretaceous lacustrine
deposits as well although the frequency of finds is rather low (POPOV 1980; JELL &DUNCAN 1986).
The dobsonflies are represented mainly by the corydalid larvae of the genus Cretachaulus
PONOMARENKO, 1976, common in the lacustrine Early Cretaceous deposits of Transbaikalia.
The Early Cretaceous aquatic beetles are represented mainly by the same families as in the Ju-
rassic, and the set of ecological groups is similar as well. They occur in different deposits, some-
times where no other aquatic insects are found. In the Late Cretaceous some most peculiar
Mesozoic taxa had disappeared completely (e.g., coptoclavids). The appearing of aquatic leaf bee-
tles of the subfamily Donaciinae in the Latest Cretaceous is noteworthy.
Main ecological events in aquatic insects history 389
The caddisflies evolved rapidly in the Early Cretaceous. Lacustrine assemblages are dominated
by several integripalpian families, first of all by extinct Vitimotauliidae, although some necrotau-
liids still survive as well. Fossil larval cases show rapid behavioral evolution over the Early Creta-
ceous. At the very end of the Early Cretaceous the diversity of caddis cases in lacustrine
assemblages fell drastically suggesting an extinction of the majority of lentic taxa. In the Late Creta-
ceous the lacustrine assemblages are extremely poor in species (usually one or two in each). On the
other hand, resin inclusions document radiation of modern families, both annulipalpian and integri-
palpian, probably mainly in running water environments.
The dominating aquatic dipteran families in the Early Cretaceous are the same as in the Jurassic.
The first appearance of aquatic brachyceran fly larvae is worth of mentioning. On the contrary, in
the Late Cretaceous few dipteran fossils occur in lacustrine deposits, in a strong contrast with di-
verse running water fauna represented in fossil resins. In particular, no chaoborids occur in lacus-
In the Jurassic and especially in the Early Cretaceous lakes the diversity of aquatic insects pos-
sessing special adaptations to swimming attracts attention. Some of them are rather unusual, like the
larvae of coptoclavid beetles (Fig. 11), and especially the nymphs of hemeroscopid dragonflies
(Fig. 12). The latter group is unique among the odonatans in its swimming adaptations. Possibly the
mayfly nymphs of the family Hexagenitidae (Fig. 13) could be referred to the nectobenthic swim-
ming fauna too. They have wide gills (tergaliae) with thickened margins, which could help the
nymphs to keep their body in water, and the larger last pair of gills could serve as oars.
In the Early Tertiary the lacustrine insect fauna for a long time remained poor and uniform, like
in the Late Cretaceous. This is a universal pattern in all types of lakes, small as well as large and
lowland as well as montane. That times the main environments of aquatic insect evolution were
likely the running waters. A recolonization of lakes was slow, and diverse lacustrine assemblages
are reappearing in the fossil record not earlier than near the Eocene/Oligocene boundary.
Fig. 11-12. Aquatic insects from the Early Cretaceous of Siberia, Russia. 11 – a specialized nectic beetle larva Coptoclava
longipoda PING 1935, Coptoclavidae; 13 – mayfly larva Protoligoneuria limai DEMOULIN 1955, Hexagenitidae, Early
Cretaceous of Brazil.
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