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Late Quaternary insects and freshwater invertebrates of the Alaskan North Slope and paleoenvironmental reconstructions in Arctic Alaska

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© INVERTEBRATE ZOOLOGY, 2019Invertebrate Zoology, 2019, 16(2): 89–125
Late Quaternary insects and freshwater invertebrates of
the Alaskan North Slope and paleoenvironmental
reconstructions in Arctic Alaska
S.A. Kuzmina1a, S.A. Elias2, A.A. Kotov3,4
1 Laboratory of Arthropods, Borissiak Palaeontological Institute, Profsoyuznaya Str. 123, 117997,
Moscow, Russia.
2 Institute of Arctic and Alpine Research, University of Colorado, Boulder, CO 80309-0450, USA.
3 Laboratory for Ecology of Aquatic Communities and Invasions, A.N. Severtsov Institute of Ecology
and Evolution, Russian Academy of Sciences, Leninsky Prospect 33, Moscow 119071, Russia.
4 Laboratory of Paleoclimatology, Kazan Federal University, Kremlevskaya Street 18, Kazan
420000, Russia.
E-mails: svkuz@yandex.ru, S.Elias@rhul.ac.uk, alexey-a-kotov@yandex.ru
ABSTRACT: A large series (26 samples) of fossil insect assemblages were excavated from
riverbank exposures at two localities at Ikpikpuk and Titaluk Rivers on the North Slope of
Alaska, U.S.A. Climatic conditions were reconstructed for the Late Pleistocene and early
Holocene based on the fossil insect assemblage data. Insects indicate the continuous
existence of a steppe-tundra community on the Alaskan North Slope during the end of the
Pleistocene and the beginning of the Holocene. The invasion of poplar during the Early
Holocene occurred within the context of the steppe-tundra community. The insect faunas
indicate plant communities dominated by grasses and other herbs, with the local presence
of tall shrubs and dwarf willows. The composition of the North Slope insect communities
during the study interval was affected by the high latitude of the localities, periglacial winds
coming off Brooks Range glaciers, and the close proximity of eolian sand and silt. The North
Slope environment differed from those in more southerly localities in Eastern Beringia,
reflecting ancient climatic and vegetational zonation.
How to cite this article: Kuzmina S.A., Elias S.A., Kotov A.A. 2019. Late Quaternary
insects and freshwater invertebrates of the Alaskan North Slope and paleoenvironmental
reconstructions in Arctic Alaska // Invert. Zool. Vol.16. No.2. P.89–125. doi: 10.15298/
invertzool.16.2.02
KEY WORDS: Pleistocene, Holocene, fossil insects, beetles, Cladocera, Beringia, steppe-
tundra.
Позднечетвертичные насекомые и пресноводные
беспозвоночные Северного Склона Аляски и
реконструкции природной обстановки для
арктической Аляски
С.А. Кузьмина1a, С.А. Элайс2, А.А. Котов3,4
1 Лаборатория артропод, Палеонтологический институт им. А.А. Борисяка, ул. Профсоюз-
ная 123, 117997, Москва, Россия.
2 Institute of Arctic and Alpine Research, University of Colorado, Boulder, CO 80309-0450, USA.
Contributions to Quaternary Zoology. Issue dedicated to the 80th anniversary of A.V. Sher.
90 S.A. Kuzmina et al.
of the modern Arctic (Colinvaux, 1967; Ager,
1982; Anderson et al., 1989), to meadow-like
tundra (Anderson, 1985), a mosaic grassland
(Schweger, 1997) to arid steppe-tundra (Sher,
1974; Matthews, 1979; Kiselev, 1981; Guthrie,
1990, 2001; Yurtsev, 2001; Zazula et al., 2003;
Kienast, 2006). These different reconstructions
are caused primarily by the different proxies
used to make them. Thus, pollen studies may
indicate tundra or meadow environments, while
plant macrofossil, mammal and insect studies
indicate steppe-tundra. The kind of environ-
mental reconstruction also depends on the re-
gion studied. For instance, in Western Beringia,
insect fossil assemblages from Late Pleistocene
cold stages (MIS4, MIS2) indicate well devel-
oped steppe-tundra in the Kolyma and Yana-
Indigirka lowlands, while tundra-like commu-
nities with little contribution from steppe spe-
cies have been identified from far northern
Introduction
The enormous ancient region known as Ber-
ingia included the northeastern part of Russia
and the northwestern regions of North America,
linked together by the Bering Land Bridge.
Beringia played a critical role in the history of
the Arctic flora and fauna (Hultén, 1937; Yurt-
sev, 1972; Hopkins, 1972; Sher, 1976; Mat-
thews et al., 2019). Because of its aridity, Ber-
ingia remained mainly unglaciated throughout
the Pleistocene. Thus Beringia was essentially
the only Arctic region that remained ice-free,
making it a refuge for Arctic flora and fauna
during Pleistocene glaciations. Beringia was
covered by different kinds of grasslands with a
minor component of trees in regions of higher
moisture. There are different opinions about the
nature of Beringian vegetation cover, ranging
from tundra-like environments similar to those
3 Лаборатория экологии водных сообществ и инвазий, Инститкт проблем экологии и
эволюции им. А.Н. Северцова РАН, Ленинский проспект 33, Москва 119071, Россия.
4 Лаборатория палеоклиматологии, Казанский (Приволжский) федеральный университет,
ул. Кремлевская 18, Казань 420000, Республика Татарстан, Россия.
E-mails: svkuz@yandex.ru, S.Elias@rhul.ac.uk, alexey-a-kotov@yandex.ru
РЕЗЮМЕ: Большая серия образцов (26 штук) с ископаемыми насекомыми была
отобрана из двух разрезов на реках Икпикпук и Титалук на Северном Склоне Аляски.
На основании изучения комплексов насекомых получены реконструкции климата
для позднего плейстоцена и раннего голоцена. Насекомые позволяют утверждать,
что ландшафты на Северном Склоне Аляски, и во время позднего плейстоцена, и в
начале голоцена, были тундростепными. Даже появление на данной территории
тополя в начале голоцена не отразилось на тундростепном сообществе насекомых.
Энтомофауна существовала в растительном сообществе из злаков, разнотравья,
локального развития среднего размера кустарников и карликовой ивы. На фауну
насекомых Северного Склона оказывали влияние северное расположение района,
стоковый ветер с хребта Брукса, где в то время было развито горное оледенение, и
близость источника эоловых песков и пыли. Природная обстановка на Северном
Склоне отличалась от того, что наблюдалось в более южных районах Восточной
Беринги, что отражает существовавшую в прошлом природно-климатическую зо-
нальность.
Как цитировать эту статью: Kuzmina S.A., Elias S.A., Kotov A.A. 2019. Late Quaternary
insects and freshwater invertebrates of the Alaskan North Slope and paleoenvironmental
reconstructions in Arctic Alaska // Invert. Zool. Vol.16. No.2. P.89–125. doi: 10.15298/
invertzool.16.2.02
КЛЮЧЕВЫЕ СЛОВА: плейстоцен, голоцен, ископаемые насекомые, жуки, Cladocera,
Берингия, тундростепь.
91Late Quaternary insects and freshwater invertebrates of the Alaskan North Slope
Siberia (Medvezhji Islands, Bolshoy Lya-
khovsky Island). Meanwhile, meadow-tundra
vegetation has been reconstructed for the south-
ern Chukotka region (Kuzmina, 2015).
The composition of vegetation zones in
Western Beringia was affected by the distance
from the Arctic Ocean, by the distance from
glaciers, and by distance from high mountain
ridges. Environmental conditions in the interior
regions in Western Beringia, such as Kolyma
Lowland, were favorable for the establishment
of steppe vegetation. Even during coldest time
of the Pleistocene, the Last Glacial Maximum
(LGM), summer temperatures were higher here
than they are today (Alfimov, Berman, 2001,
Alfimov et al., 2003). In contrast, temperature
reconstructions made from insect fossil assem-
blages in Eastern Beringia (Elias, 2001) indicat-
ed that summer temperatures during the LGM
were lower than today.
A paleontological expedition to the North
Slope of Alaska took place in June 2003 after
the 3rd International Mammoth Conference in
Dawson City and Whitehorse, Yukon. Paul
Matheus was a member of the conference host
committee; Andrei Sher was one of distinguished
guests and a session chair. During a pre-confer-
ence meeting, Paul and Andrei discussed possi-
bility of joint fieldwork in Alaska. It was diffi-
cult bureaucratic task since the Russian mem-
bers (A.V. Sher and S.A. Kuzmina) needed
visas to enter Canada and USA; combining the
conference with a field trip to Alaska provided
a good opportunity to obtain these visas. A.V.
Sher wanted to visit two sites to collect insect
fossils: the Alaskan analogue of Siberian “Ye-
doma” on the North Slope of Alaska and a
tephra buried soil on Seward Peninsula. Insect
remains had already been observed at both sites
(Nelson, Carter, 1987; Goetcheus et al., 1994,
2001; Goetcheus, 2001) but the sections were
tested differently than the methods used for
Siberian sites. Thus Sher’s idea was to apply the
“Russian” method of insect sampling and subse-
quent treatment, involving the screening of large
volumes of the sediment in the field, and the
sampling of entire sections at exposures (not
just the most interesting layers); and, most im-
portantly, the identification of the Alaskan in-
sect remains by a Russian expert. Such method-
ological comparisons would be helpful in the
understanding of the similarities and differenc-
es between Western and Eastern Beringia insect
faunas.
The plan worked well. A.V. Sher, S.A.
Kuzmina and P. Matheus collected rich fossil
insect assemblages from two sites on the North
Slope and one site on Seward Peninsula (Kuzmi-
na, Elias, 2019). The results of the expedition to
Seward Peninsula have been published (Kuzmi-
na et al., 2008) but the results of the North Slope
investigations have not been published until
now. The authors have taken the opportunity to
complete this important research that began
with Andrei Sher’s 2003 initiative.
Regional setting
1. Modern environment and climate
The North Slope of Alaska (Fig. 1) is situat-
ed between the Brooks Range and the Arctic
Ocean. Today it is a treeless region of flat plains
and gentle foothills (Fig. 2 A, C, D). While the
North Slope itself has never been glaciated, the
Brooks Range was covered by ice during LGM
(Kaufman, Manley, 2004; Briner, Kaufman,
2008). These mountains contribute to windy
weather on the plain. During the cold stages of
the Late Pleistocene, dry winds coming off the
Brooks Range helped shape the Ikpikpuk Sand
Sea with its famous sand dunes (Carter, 1981).
These winds also helped to accumulate loess
deposits exposed along the Ikpikpuk and Ti-
taluk rivers. A significant part of the Pleistocene
deposits in the study area were formed by allu-
vium derived from numerous streams flowing
out of the Brooks Range. The studied sections of
these two rivers contain gravel and sand units
with cross bedding (Fig. 3).
The recent climate of the interior regions of
the North Slope is cold and relatively dry. The
mean annual air temperature is –10.2 °C, the
warmest month is July (12–13 °C), and mean
annual precipitation is 122 mm, mostly falling
as snow (Kene et al., 2014). The active layers
above permafrost are 15–40 cm thick in summer
92 S.A. Kuzmina et al.
Fig. 1. Map of the study area.
Рис. 1. Карта изученного района.
Fig. 2. Landscape and vegetation of North Slope tundra in early June 2003.
A — aerial view of the Ikpikpuk River with a river bluff, sand bars, lakes and snow; B — upland vegetation includes
short willows, grasses, and sedges; the surface is covered by sand from a nearby outcrop; C — field camp on sand bar
of the Titaluk River, showing belts of tall willows, boggy meadow, and shrub alder; D — spring water fills old channels
in the floodplain.
Рис. 2. Ландшафт и растительность тундры Северного Склона в начале июня 2003 г.
A — вид из вертолета на реку Икпикпук, речные обрывы, песчаные отмели, озера и снег; B — растительность
тундры вне долины реки с низкими ивами, травами и осоками, почва покрыта песком, надутым с ближайшего
обрыва; C — полевой лагерь на песчаном пляже реки Титалук, видны полосы высокой кустарниковой
растительности из ивы и ольховника, и заболоченные луга; D — весенняя вода заполнила ложбины в пойме реки.
93Late Quaternary insects and freshwater invertebrates of the Alaskan North Slope
Fig. 3. Ikpikpuk Section “Little Supreme Bluff” (LSB).
Рис. 3. Разрезмаленький верховой обрыв” (ЛСБ) на реке Икпикпук.
(Bockheim, Hinkel, 2005), keeping water on
the surface (Fig. 2 D) despite the low precipita-
tion. Modern vegetation includes sedge-grass
and moss tundra with dwarf shrubs. Middle
sized willow and alder shrubs occur in parts of
the river valleys (Fig. 2 C). Balsam poplar
(Populus balsamifera L.) may grow in valleys
(Bockheim et al., 2003) but these trees are
absent near the studied sections.
2. Stratigraphic framework
The studied sections are located on the Ik-
pikpuk River and its tributary, the Titaluk River.
The river sources lie in the foothills of the
Brooks Range. Upstream channels contain Cre-
taceous sandstone bedrock; the eroded sand-
stone provides coarse and fine sand, and gravel
for the river alluvium. The sands yield small
pieces of amber and coal, probably of Creta-
ceous origin.
The Ikpikpuk sand sea (Carter, 1981), situ-
ated near confluence of the Ikpikpuk and Ti-
taluk Rivers, deposited sand more-or-less con-
tinuously in and around the two rivers during the
Late Pleistocene. The dune field was active
between 41 and 16.7 cal ka BP, then between
12.9 and 8.7 cal ka BP (Carter, 1993). Wind
carried sand from the dune deposit to the studied
sections. When the dune field was active, the
local vegetation was constantly being covered
by sand; mosses could not survive here, and the
land surface became unusually dry (Fig. 2 B).
Such xeric places increased the biotope diversi-
ty of the study area. Modern ground beetles
(Carabidae) collected from interior regions of
the North Slope (Nelson, 2001) include 56
species while only 9 species have been found at
coastal sites. Our collection of modern beetles
includes such common Pleistocene species as
Lepidophorus lineaticollis Kby, 1837, Poecilus
nearcticus Lindr., 1966, Stereocerus haemato-
pus (Dej., 1831), and Amara alpina Payk.,
1790. These beetles were found at the edge of
the river bluff on sandy soil.
94 S.A. Kuzmina et al.
Table 1. Radiocarbon data from the key sections of the North Slope.
Таблица 1. Радиоуглеродные датировки из ключевых разрезов Северного Склона.
Both rivers are surrounded by vast sand bars
(Fig. 2 A, C) where rich Pleistocene bone as-
semblages have been collected (Guthrie, Stok-
er, 1990; Mann et al., 2010, 2013). The Pleis-
tocene megafauna of the North Slope was dom-
inated by mostly horses, with Pleistocene bison,
caribou, woolly mammoth, and muskox, with
rare specimens of moose, saiga antelope, mast-
odon, and predators including short-faced bear,
cave lion, and wolf (Mann et al., 2010, 2013).
Most of the Pleistocene sections of the North
Slope are situated along rivers. The key sections
are Itkillik (Kanevskiy et al., 2011, 2016;
Lapointe et al., 2017), Nigu (Mann et al., 2010),
Etivluk (Mann et al, 2002; Gaglioti et al., 2017)
Titaluk and Ikpikpuk (Mann et al., 2010, 2013,
2015; Gaglioti et al., 2017, 2018) and a sedi-
ment core from Burial lake (Finkenbinder et al.,
2015). The river outcrops (except buried sand
dunes such as Carter’s section at the Ikpikpuk
From: Nelson, Carter, 1987 (a); Mann et al., 2010 (b); Lapointe et al., 2017 (c); Kanevskiy et al., 2011 (d); Trimble,
Robinson, 1989 (e); Mann et al., 2002 (f).
Unit 14C age
site Ikpikpuk Titaluk Nigu Etivluk Itkillik
2310±70 b 2500±50 e 4900±60 f 5320 ± 35 d
5430±60 b 8610 ± 35 d
Holo-
cene
6070±60 b
8710±140 a 8080±50 b 8360±70 f
9050±60 b 8870±60 f
9250±70 b 9020±50 f
9340±60 b 9450±60 b 9570±60 f
9480±50 b 9510±60 b 9670±50 b
9540±160 a 10,920±50 b
D
Populus
Bed
10,190±80 b
14,370±100 c
16,550±75 d
19,950±180 c
24,290±190 c
25,450±280 c
26,300±130 d
28,330±390 c
C
Yedoma
41,700±460 d
10,230±160 a 10,470±50 f
11,860±50 b 11,370±50 b 12,450±60 f
13,670±130 a 12,230±50 b 13,420±50 f
B upper
sand
17,730±110 a
20,360±190 a
24,740±32 a 28,000± 240 e 29,570±150 f
B middle
sand
30,260±530 a 34,920±260 f
B lower
sand
>39,000 a 43,000± 1400 e 47,830±
1300 f
>40,000 a
>42,500 b
>47,770 b
A gravel >49,000 a
95Late Quaternary insects and freshwater invertebrates of the Alaskan North Slope
River) usually consist of fluvial deposits: gravel
and sand. In some sections (Itkillik, Titaluk) the
uppermost unit consists of silts with ice wedges
that are similar to the Siberian “Yedoma” de-
posits (Kanevskiy et al., 2016). Another inter-
esting feature is the “Populus Bed” (Nelson,
Carter, 1987). There are depressions in the main
Pleistocene unit filled by early Holocene sedi-
ments containing well-preserved leaves, logs
and stumps of Populus balsamifera. Since the
modern limit of this tree species is south of the
North Slope, the presence of fossil poplar indi-
cates warmer than present climate during the
early Holocene.
3. Radiocarbon dating
Radiocarbon dates were obtained from sev-
eral sections (Table 1). The best studied are the
sections at Ikpikpuk River (Little Supreme Bluff
and Cottonwood Bluff) where the main focus
was dating of the Populus Bed. Lower gravel
unit at Ikpikpuk River is older than 50 ka BP
(infinite radiocarbon ages). The same unit was
observed at the base of the Titaluk sections. The
main sand body was accumulated from ca. 40 ka
or earlier, to 10–11 ka BP. The Yedoma-ana-
logue unit was observed at the Tituluk site, and
was well studied at the Itkillik site, but was
absent from the Ikpikpuk site. It is likely that the
sandy unit of the Ikpikpuk section yielded a
nondepositional unconformity about 30 ka BP.
The age of the Titaluk Yedoma-analogue unit
remains uncertain but stratigraphic correlation
with the Itkillik section allows us to infer that it
formed during LGM.
4. Studied sections
Fossil insects were collected from two sec-
tions (Fig. 1): 1 — Ikpikpuk River, Little Su-
preme Bluff (LSB) (69°3521 N, 154°5614
W) and Titaluk River, Russian Section (RS)
(69°2548N, 155°926.2 W). The Carter Sec-
tion at the Titaluk River (Gaglioti et al., 2018)
was visited but useful layers for insect sampling
were not found. Since the expedition took place
in early June, the water level in the rivers was
high. Only the lower part of the LSB exposure
was sampled (Fig. 3); RS (Figs 4, 5) was sam-
pled from all accessible units, including two
Holocene buried channels with poplar leaves.
Section LSB at the Ikpikpuk River is one of
the best-studied in the area (Mann et al., 2002,
2010; Gaglioti et al., 2017). Our team visited
the exposure first and planned to stay there for
several days but the section was poorly exposed,
as the top was covered by snow (Fig. 3). We
tested only lower gravel unit, the Purple Gravel
of Mann et al. (2010). The unit consists of cross-
beds of iron-stained pebbly gravel with plant
debris and willow roots. A radiocarbon age of
the unit (from willow wood) is infinite — more
than 47 ka yr.
The Russian Section (RS) at the Titaluk
River had not been previously described. This
section was better exposed (Figs 4, 5); we spent
about two weeks working there. The lower part
of the section is similar to LSB. There is cross-
bedded gravel with plant debris; the unit is
traced along whole outcrop, about 1 km long.
The deposit is exposed from the river level up to
2–2.5 m.
The gravel unit is overlaid with a visible
unconformity by a sand unit. It consists of cross-
beds and horizontal sand with numerous thin
layers of plant debris. The plant layers were
found to contain abundant fossil insect remains.
The age of the unit is uncertain. Sher considered
that the unit could be very old, either from the
middle or early Pleistocene. Infinite radiocar-
bon ages from a similar sand deposit at the
Cottonwood Bend Section at Ikpikpuk River
(Nelson, Carter, 1987) support this assumption.
However, other radiocarbon dates from 30 to 11
ka yr (Table 1) indicate that the whole unit
probably belongs to the late Pleistocene. Per-
haps the lower part of the sand is older.
Ice-rich silt with ice wedges are situated at
the top of the section (Fig. 5). This unit is very
similar to Siberian Yedoma, based on the field
experience of Sher and Kuzmina; the sediment
was wet because of melting, and plant inclu-
sions were not evident. In contrast to the lower
units, the silt looked fossil-free. We paid special
attention to the insect sampling of the silt. Large
volume screening of sediments yielded good
insect assemblages from this organic-poor sed-
96 S.A. Kuzmina et al.
Fig. 4. Titaluk “Russian Section” (RS), profiles 1 (lower part) and 4.
Рис. 4. РазрезРусский” (РС) на реке Титалук, расчистки 1 (низ) и 4.
iment. Yedoma-like sediments were observed
in one profile only. This unit was inserted into
the sand and probably filled an eroded depres-
sion. As we know from the well-developed
Itkillik Yedoma deposit, the age of the ice- rich
silt could range any time from 42 to 14 ka. We
have to take into consideration that Yedoma
deposits are quite rare on the Alaskan North
Slope, in contrast to northeastern Siberia where
they are widespread. On the North Slope, loess
accumulation was local and was probably con-
current with more common sand dunes, and
fluvial deposition.
Populus Beds were found in two profiles at
RS (Figs 4, 5). These sediments are represented
by fine sand and abundant plant layers consist-
ing of well-preserved poplar leaves, small twigs,
and wood pieces. Populus Beds filled eroded
cuts inside the sandy unit. The age range of the
Populus beds is 10–8 ka yr. (Table 1). This
interval includes two episodes of warming: pre-
and post-Younger Dryas observed in both East-
ern Beringia (Ager, 1982; Edwards, Barker,
1994; Mann et al., 2002, 2010) and Western
Beringia (MacDonald et al., 2000; Smith et al.,
2004; Grosse et al., 2007).
The second warming (around 9–8 ka), which
corresponds with Boreal Period of the Blytt-
Sernander scheme, was probably the warmest
interval of the Holocene in Beringia (Kaufman et
al., 2004). Trees spread north to coastal areas in
Siberia (Kaplina, Lozhkin, 1982; Kuzmina, Sher,
2006), spruce forest reached the central Brooks
Range (Anderson, Brubaker, 1994), and beaver
97Late Quaternary insects and freshwater invertebrates of the Alaskan North Slope
Fig. 5. Titaluk “Russian Section” (RS), profiles 1 (upper part) and 6.
Рис. 5. РазрезРусский” (РС) на реке Титалук, расчистки 1 (верх) и 6.
98 S.A. Kuzmina et al.
Fig. 6. Scheme of ecological group variation in Ikpikpuk and Titaluk fossil insect assemblages.
Рис. 6. Схема изменений соотношения экологических групп в комплексах ископаемых насекомых с
рек Икпикпук и Титалук.
99Late Quaternary insects and freshwater invertebrates of the Alaskan North Slope
occupied formerly treeless landscapes in Alaska
(Robinson et al., 2007). Climate on the North
Slope was about 2 to 3°C warmer than today;
and moisture was lower (Nelson, Carter, 1987).
Material and Methods
Insect samples were taken from 9 profiles
(Figs 2–5), including 5 samples from the gravel
unit at Ikpikpuk, 4 samples from the lower
gravel unit similar to the lower unit at Titaluk, 7
samples from the main sandy unit at Titaluk, 3
samples from the Yedoma-analogue deposit,
and 6 from the Populus Beds at Titaluk. The
volume of each sample was different. Only 1–2
kg of sediment was sampled from the plant
debris beds with visible insect remains; from
less organic-rich deposits we took up to 50 kg
per sample. Insects were concentrated by wet
screening through 0.4 mm mesh and dried in the
field, followed by later sieving of this concen-
trate in the laboratory, and picking of fossil
insects with a binocular microscope (Sher,
Kuzmina, 2007; Kuzmina et al., 2008).
Ecological interpretation of the fossil insect
follows the previous studies (Matthews, 1983;
Kuzmina et al., 2008, 2014). The method is
based on the analysis of the proportion of eco-
logical groups present in a particular sample.
Ecological groups (Table 2, Fig. 6) include:
1. st — indicators of Pleistocene steppe-
tundra environments such as Aphodius pectora-
lis LeC. 1857, Morychus aff. aeneolus (LeC.,
1863), Stephanocleonus confusus And., 1987,
Coniocleonus parshus (And., 1987). These spe-
cies live today in relict steppe communities in
Alaska and the Yukon, or in xeric habitats to the
south of this region.
The pill beetle Morychus aff. aeneolus has
previously been recorded by other workers as
Morychus sp. A, Morychus sp. B, or Morychus
sp. (Matthews, 1968, 1983; Matthews, Telka,
1997). The taxonomic status of these very com-
mon Pleistocene beetles remains uncertain. We
found a number of modern Morychus speci-
mens on a relict steppe site near Kluane Lake,
Yukon. These modern specimens are similar to
the Pleistocene fossils in that they have a small
elytral shoulder and shortened wings. At the
same time the normal full wing M. aeneolus
individuals are found on the Kluane steppe. The
male genitalia of these two morphological types
are slightly different. Since specimens of M.
aeneolus are quite variable in shape, including
shoulder size (Johnson, 1991), we can not con-
fidently describe our specimens as a new spe-
cies. All the Pleistocene specimens we found
belong to the small shoulder morph and, if the
species is indeed M. aeneolus, it is represented
by stable (more than one million years) short-
wing variation. So we decided to identify the
Pleistocene beetles as M. aff. aeneolus (Kuzmi-
na, Matthews, 2012; Murton et al., 2017).
Following the revision of Anderson (1987,
1989), Coniocleonus Motsch., 1860 and Steph-
anocleonus Motsch., 1860 are regarded here as
subjective synonyms. Anderson (1987) exam-
ined type species of two genera and made a
conclusion: “In view of this variation and the
lack of other distinguishing characters, I cannot
accept other than that Stephanocleonus is sim-
ply an apterous Coniocleonus. I do not think the
rostral differences between the type species
warrant separate generic status.” (Anderson,
1987, p.453). “I choose to give the name Steph-
anocleonus priority over Coniocleonus because
the former has been the only name used in
reference to the Nearctic species here placed in
the genus. Furthermore, no major works have
recently appeared dealing with Coniocleonus,
however, a recent review by Ter-Minasyan
(1979) of the Stephanocleonus of the Palearctic
Region includes species that in my opinion are
congeneric with species in North America”
(Anderson, 1987, p.454).
This decision was accepted (or partly ac-
cepted) by the international entomological com-
munity (for example Alonso-Zarazaga, 1999
noticed only Palearctic Coniocleonus) but was
very inconvenient for environment reconstruc-
tion in Beringia. We can easily recognize the
remains of two genera in the fossil assemblages.
Coniocleonus sensu lato indicates meadow-
steppe or dry tundra environments, but Stepha-
nocleonus sensu stricto indicates thermophil-
ous steppes. Fortunately, the genus Coniocleo-
100 S.A. Kuzmina et al.
sample eco
Ikpikpuk B-0- Titaluk B-0-
taxon code
1 2 4 4a 5 1 2 3 4
Ord. Coleoptera unit A A A A A A A B B
Fam. Carabidae
Subfam. Nebriinae
Notiophilus aquaticus (L., 1758) dt _ _ _ _ _ _ _ 1 _
Notiophilus sp. dt _ _ _ _ _ _ _ _ _
Subfam. Carabinae
Carabus truncaticollis Esch., 1833 mt _ _ _ _ _ _ _ _ _
Carabus sp. mt _ _ _ 1 _ 1 _ 1 _
Subfam. Elaphrinae
Elaphrus angusticollis Sahlb., 1844 r&a _ _ _ _ _ _ _ _ _
E. clairvillei Kby., 1837 r&a _ _ _ _ _ _ _ _ _
Subfam. Scaritinae
Dyschirius nigricornis Motsch.,
1844 dt _ _ _ _ _ _ _ _ 1
Subfam. Trechinae
Asaphidion alaskanum Wick., 1919 r&a _ _ 1 _ _ _ _ _ _
A. yukonense Wick., 1919 r&a _ _ _ 1 _ _ _ _ _
Bembidion (Asioperyphus)
bimaculatum Kby., 1837 r&a
_ _ _ 2 1 1 _ _ _
B. (Asioperyphus) sordidum (Kby.,
1837) r&a _ _ _ _ _ _ _ _ _
B. (Asioperyphus) umiatense
Lindr., 1963 r&a _ _ 2 2 _ _ _ 1 _
B. (Peryphanes) dauricum
(Motsch., 1844) dt _ _ _ _ _ _ _ _ _
B. (Peryphanes) spp. dt _ _ _ _ _ 1 _ _ _
B. (Plataphus) hyperboraeorum
Mnst., 1923 r&a _ _ 1 _ _ _ _ _ _
B. (Plataphodes) arcticum Lindr.,
1963 r&a _ _ _ _ _ _ _ _ _
Bembidion sp. r&a _ _ _ _ _ _ _ _ _
Subfam. Harpalinae
Tribe Harpalini
Dicheirotrichus mannerheimi
(Sahlb., 1844) dt _ _ _ 2 _ _ _ _ _
Harpalus amputatus Say, 1830 dt _ _ _ 1 _ _ _ _ _
H. vittatus alaskensis Lindt., 1968 dt _ _ _ 2 2 _ _ _ _
Harpalus sp. dt _ _ _ _ _ 2 _ _ _
Tribe Lebiini
Cymindis unicolor Kby., 1837 dt _ _ _ _ _ _ _ _ _
Tribe Platynini
Agonum (Europhilus) exaratum
(Mnnh., 1853) r&a
_ _ _ _ _ _ _ _ _
101Late Quaternary insects and freshwater invertebrates of the Alaskan North Slope
Table 2. List of insects from Ikpikpuk and Titaluk sections.
Таблица 2. Список насекомых из разрезов Икпикпук и Титалук.
5 6 7 7a 8a 8b 9 11 11a 12 13 13a 14 15 16 17 18
B B B B C C C D D D A A A B D D D
_ _ _ _ _ _ _ _ _ _ 1 1 _ 2 _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ 1 _ _ _ _
_ _ _ 2 1 _ _ _ _ _ _ _ _ _ _ 2 _
_ _ _ _ _ _ _ _ _ _ 1 1 _ 1 1 _ _
_ _ _ _ _ _ _ _ _ _ 1 _ _ _ _ 1 1
_ _ _ _ _ _ _ _ _ _ _ 1 _ _ _ _ _
_ _ _ _ _ _ _ _ _ _ _ 1 _ 2 _ _ _
_ _ _ _ _ _ _ _ _ _ 1 _ _ _ _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
_ _ _ 1 _ _ _ _ _ _ 1 1 _ _ _ _ _
_ _ _ _ _ _ _ _ _ _ 2 1 _ 1 _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ _ 2 _ _ _
_ _ _ _ _ _ _ _ _ _ 3 _ _ _ _ _ _
_ _ 1 _ _ _ _ _ _ _ _ _ _ _ _ _ _
_ _ _ 1 _ _ _ _ _ _ 1 1 _ _ _ _ _
_ 1 _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
_ _ _ _ 1 _ _ _ _ _ 1 _ _ _ _ _ _
_ _ _ _ 1 _ _ _ _ _ _ _ _ _ _ _ _
_ 2 _ 11 1 2 _ _ _ 2 1 1 1 _ _ 1 _
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ 1 _ _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ _ 1 _ _ _
102 S.A. Kuzmina et al.
sample eco
Ikpikpuk B-0- Titaluk B-0-
taxon code
1 2 4 4a 5 1 2 3 4
Tribe Pterostichini
Poecilus (Derus) nearcticus Lindr.,
1966 dt _ 1 1 3 2 _ _ 2 _
Pterostichus (Cryobius) arcticola
Chad., 1868 mt
_ _ _ 7 18 14 14 7 _
P. (Cryobius) brevicornis (Kby.,
1837) mt _ _ 18 21 29 5 5 31 _
P. (Cryobius) caribou Ball, 1962 mt _ _ 6 _ 7 1 _ 1 _
P. (Cryobius) empetricola (Dej.,
1828) mt _ _ 1 _ _ _ _ _ _
P. (Cryobius) kotzebuei Ball, 1962
? mt _ _ _ _ _ _ _ _ _
P. (Cryobius) nivalis Sahlb., 1844 dt _ _ 29 _ 1 _ _ 5 _
P. (Cryobius) parasimilis Ball,
1962 mt _ _ _ _ 3 4 4 3 _
P. (Cryobius) pinguedineus Esch.,
1823 mt _ 1 7 19 12 4 2 11 1
P. (Cryobius) pinguedineus group mt _ _ _ _ _ _ _ _ _
P. (Cryobius) similis Mann., 1852 dt _ _ 6 17 23 13 7 11 1
P. (Cryobius) soperi Ball, 1966 mt? _ _ _ _ _ 5 _ _ _
P. (Cryobius) tareumiut Ball, 1962 mt _ _ _ 35 19 8 3 12 2
P. (Cryobius) ventricosus Esch.,
1823 mt _ _ 9 12 28 4 3 7 1
P. (Cryobius) spp. mt 1 _ 15 _ _ 17 5 1 2
P. (Lenapterus) agonus Horn., 1880 mt _ _ 1 _ _ _ _ _ _
P. (Lenapterus) costatus (Men.,
1851) mt _ _ _ 2 1 _ _ _ _
P. (Lenapterus) vermiculosus
(Men., 1851) mt _ _ _ 1 _ _ _ _ _
P. (Steropus) circulosus Lindr.,
1966 ? dt _ _ _ _ _ _ _ _ _
P. (Tundraphilus) sublaevis
(Sahlb., 1880) dt _ _ 2 5 7 1 _ 2 1
Stereocerus haematopus (Dej.,
1831) dt _ 1 1 7 4 4 _ 2 _
Tribe Zabrini
Amara (Amarocelia) erratica Duft.,
1812 dt _ _ _ _ _ _ _ _ _
A. (Amarocelia) interstitialis Dej.,
1828 dt _ _ 1 _ _ _ _ _ _
A. (Bradytus) glacialis Mnnh., 1853 dt _ _ _ _ _ _ _ _ _
A. (Curtonotus) alpina Payk., 1790 dt _ _ 7 19 32 63 27 41 3
A. (Curtonotus) bokori Csiki, 1929 dt _ _ _ 1 _ _ _ _ _
A. (Paracelia) quenseli Schnh.,
1806 dt _ _ _ _ _ _ _ _ _
Fam. Dytiscidae
Subfam. Agabinae
Agabus confinis (Gyll., 1808)? r&a _ _ _ _ _ _ _ _ _
103Late Quaternary insects and freshwater invertebrates of the Alaskan North Slope
Table 2 (continued).
Таблица 2 (продолжение).
5 6 7 7a 8a 8b 9 11 11a 12 13 13a 14 15 16 17 18
_ 4 1 25 2 2 _ 1 1 1 1 2 _ 2 _ 5 3
_ 16 3 54 13 10 _ 1 2 3 53 35 56 54 13 10 7
_ 19 7 75 11 12 3 _ 2 3 28 19 29 39 19 11 3
_ _ _ 13 4 _ _ _ _ _ 3 9 9 17 1 _ _
_ _ _ 2 _ _ _ _ _ _ _ _ _ _ _ _ _
_ 3 _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
_ _ _ 1 _ _ _ _ _ _ _ _ _ _ _ _ _
_ 5 _ 27 _ _ _ _ _ 1 _ _ 10 17 2 2 _
_ 28 5 166 14 8 3 4 3 7 26 31 51 43 22 21 7
_ _ _ 30 6 1 _ _ _ _ 8 4 _ 2 _ _ _
_ 14 _ 41 6 3 _ _ _ _ 8 7 5 6 2 1 1
_ _ _ 4 _ _ _ _ _ _ _ _ 5 2 _ 2 _
_ 13 4 232 12 13 1 _ 4 7 21 22 55 34 11 8 2
_ 7 _ 99 6 2 _ 1 _ 3 16 7 14 20 4 3 _
_ 8 3 39 _ _ _ 2 _ 6 13 13 1 19 3 1 _
_ _ _ 2 _ _ _ _ _ _ 1 _ _ 3 _ _ _
_ _ _ 2 _ _ _ _ _ 2 _ _ _ _ 1 1 _
_ _ 1 _ _ _ _ _ _ _ _ 2 _ 1 _ _ _
_ _ _ _ _ _ _ _ _ _ 1 _ _ _ _ _ _
_ 6 _ 42 2 1 1 1 _ 6 4 6 4 2 1 2 1
_ 4 _ 22 5 4 _ 1 _ 2 3 6 3 5 4 4 2
_ _ _ _ _ _ _ _ _ _ 1 _ _ _ _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
_ _ _ 1 _ _ _ _ _ _ _ _ _ _ _ _ _
1 65 25 652 55 37 3 2 1 21 66 54 154 137 50 48 32
_ _ 2 _ 1 1 _ _ _ _ _ 5 6 2 _ 1 _
_ _ _ _ _ _ _ _ _ _ _ _ _ 1 _ _ _
_ _ _ _ _ _ _ _ _ _ 5 _ _ _ _ _ _
104 S.A. Kuzmina et al.
sample eco
Ikpikpuk B-0- Titaluk B-0-
taxon code
1 2 4 4a 5 1 2 3 4
A. moestus (Curt., 1835) r&a _ _ _ _ 2 2 3 2 _
A. serricornis (Payk., 1799) ? r&a _ _ _ _ _ _ _ _ _
A. thomsoni (Sahlb., 1871) r&a _ _ _ _ _ _ _ _ _
Agabus spp. r&a _ _ _ _ _ _ _ _ _
Ilybius picipes (Kby., 1837) r&a _ _ _ _ _ _ _ _ _
Subfam. Colymbetinae
Colymbetes dahuricus Aub., 1837 r&a _ _ _ _ _ _ _ _ _
C. dolabratus (Payk., 1837) r&a _ _ _ _ 1 _ _ _ _
Colymbetes sp. r&a _ _ _ _ _ _ _ _ _
Subfam. Hydroporinae
Hydroporus lapponum (Gyll.,
1808) r&a _ _ _ _ _ _ _ _ _
H. morio Aub., 1838 r&a _ _ _ _ 1 _ _ _ _
H. polaris Fall, 1923 r&a _ _ _ _ _ 2 _ 1 _
H. striola (Gyll., 1826)? r&a _ _ _ _ _ _ 1 _ _
Hydroporus spp. r&a _ _ _ _ _ _ _ _ 1
Oreodytes laevis (Kby., 1837) r&a _ _ _ _ _ _ _ _ _
O. recticollis (Fall, 1926) r&a _ _ _ _ _ _ _ _ _
O. sanmarkii (Sahl., 1826) r&a _ _ _ 1 _ _ _ _ _
Fam. Hydrophilidae
Subfam. Helophorinae
Helophorus sempervarians Ang.,
1970? r&a _ _ _ _ _ _ _ _ _
H. splendidus Sahlb., 1880 r&a _ _ _ 1 _ 2 _ 2 _
Helophorus sp. r&a _ _ _ _ _ _ _ _ _
Subfam. Hydrophilinae
Cymbiodyta sp. r&a _ _ _ _ _ _ _ _ _
Hydrobius fuscipes L., 1758 r&a _ _ _ _ _ _ _ _ _
Subfam. Sphaeridiinae
Cercyon sp. pl _ _ _ _ _ _ _ 1 _
Fam. Leiodidae
Subfam. Leiodinae
Agathidium angulare Mann., 1852 pl _ _ _ _ _ _ _ _ _
Hydnobius sp. pl _ _ _ _ _ _ _ _ _
Leiodes collaris (LeC., 1850) pl _ _ _ 1 _ _ _ _ _
Subfam. Cholevinae
Catops basilaris Say, 1823? pl _ _ _ _ _ _ _ 1 _
Fam. Silphidae
Subfam. Silphinae
Thanatophilus lapponicus (Hbst.,
1793) _ _ _ 1 _ _ _ _ _
105Late Quaternary insects and freshwater invertebrates of the Alaskan North Slope
Table 2 (continued).
Таблица 2 (продолжение).
5 6 7 7a 8a 8b 9 11 11a 12 13 13a 14 15 16 17 18
_ 2 _ _ _ _ _ _ 3 _ _ 1 _ 1 1 1 _
_ _ _ _ _ _ _ 8 _ 4 _ _ _ _ _ _ _
_ _ _ 6 _ 1 _ _ _ _ _ _ _ _ _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ 5 1 _ _ 1
_ _ _ _ _ _ _ _ _ _ _ 1 _ _ _ _ _
_ _ _ _ _ _ _ _ 3 _ _ _ _ _ _ _ _
_ _ _ 1 1 _ _ _ _ _ 1 2 _ _ _ _ 2
_ _ _ _ _ _ _ _ _ _ 1 _ _ _ _ _ _
_ _ _ 1 _ 2 _ _ _ _ 1 _ _ _ _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
_ _ 2 3 1 _ _ _ 2 _ 3 4 1 4 _ 1 _
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
_ _ _ _ _ _ _ 1 _ _ _ 2 _ _ _ _
_ _ _ _ _ _ _ _ _ 1 _ _ _ _ _ _ _
_ _ _ _ _ _ _ _ _ _ _ 1 _ _ _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ _ 1 _ _ _
_ 1 1 2 1 1 _ _ 1 1 2 _ 2 1 _ _ _
_ _ _ _ _ _ _ _ _ _ _ 1 _ _ _ 1 _
_ _ _ _ _ _ _ _ _ _ _ _ _ 1 _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ 1 1 _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
_ _ _ _ _ _ _ _ _ _ 1 1 1 2 _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 1 _
_ 1 _ 1 _ 1 _ _ _ _ _ 1 _ _ _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
106 S.A. Kuzmina et al.
sample eco
Ikpikpuk B-0- Titaluk B-0-
taxon code
1 2 4 4a 5 1 2 3 4
Th. sagax (Mann., 1853) _ _ _ _ _ _ _ _ _
Fam. Staphylinidae
Subfam. Omaliinae
Tribe Anthophagini
Acidota quadrata (Zett., 1840) pl _ _ _ _ _ _ _ _ _
Eucnecosum sp.? pl _ _ _ _ _ 1 _ _ _
Olophrum latum Maekl., 1853 r&a _ _ _ _ _ _ _ _ _
O. rotundicolle (Sahlb., 1830) r&a _ _ _ _ _ _ _ _ 1
Olophrum sp. r&a _ _ 1 _ _ _ _ _ _
Tribe Omaliini
Micralymma brevilingue Schtd.,
1845 dt _ _ _ _ _ 2 1 1 _
Subfam. Tachyporinae
Tribe Tachiporini
Tachinus apterus Maekl., 1853 ar _ _ _ _ _ _ _ _ _
T. brevipennis Sahlb., 1880 mt _ _ _ 4 _ 35 11 5 5
T. instabilis Maekl., 1853 pl _ _ _ _ _ _ _ 1 _
Subfam. Steninae
Stenus sp. r&a _ _ _ 1 _ _ _ _ _
Subfam. Paederinae
Lathrobium nigrum LeC., 863 pl _ _ _ _ _ _ _ _ _
L. sibiricum Fauvel, 1875 pl _ _ _ _ _ _ _ 1 _
L. washingtoni Csy., 1905 pl _ _ _ 2 _ 1 _ _ _
Subfam. Staphylininae
Philonthus subvirescens Thom.,
1884 pl _ _ _ _ _ _ _ _ _
Quedius fellmani (Zett., 1838) pl _ _ _ _ _ _ _ _ _
Q. sublimbatus Maekl., 1853? pl _ _ _ _ _ _ _ _ 1
Quedius sp. pl _ _ _ 1 _ _ _ _ _
Fam. Scarabaeidae
Aphodius pectoralis LeC., 1857? st _ _ _ _ _ _ _ _ _
Aphodius sp. pl _ _ _ _ _ _ _ _ _
Fam. Byrrhidae
Subfam. Byrrhinae
Morychus aff. aeneolus (LeC.,
1863) st _ _ 10 29 16 _ _ 5 2
Simplocaria sp. dt _ _ _ _ _ _ _ _ _
Subfam. Syncalyptinae
Curimopsis albonotata (LeC.,
1861) dt _ _ _ 1 3 1 _ _ _
C. echinata (LeC., 1850) dt _ _ 1 _ _ _ _ _ 1
C. moosilauke Johns., 1986 dt _ _ _ 2 _ _ _ 1 _
107Late Quaternary insects and freshwater invertebrates of the Alaskan North Slope
Table 2 (continued).
Таблица 2 (продолжение).
5 6 7 7a 8a 8b 9 11 11a 12 13 13a 14 15 16 17 18
_ _ _ _ _ _ _ _ _ _ _ _ 1 1 _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ 1 _ _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 1
_ _ _ _ _ _ _ _ _ _ _ 3 _ 3 _ 1 _
_ _ _ _ _ _ _ _ _ _ 1 _ _ 1 _ _ _
_ 1 3 2 2 _ _ 1 _ _ _ _ _ 2 _ _ _
_ _ _ _ _ _ _ _ _ _ 1 _ _ _ _ _ _
_ 14 9 14 8 9 1 1 1 3 8 4 38 20 _ 5 1
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ 1 _ _ _ _
_ _ _ _ _ _ _ _ _ _ _ 2 1 _ _ _ _
_ 1 _ _ 1 _ _ _ _ _ 1 _ _ 1 _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
_ _ _ _ _ _ _ _ _ 1 1 _ _ _ _ _ 1
_ _ _ _ _ _ _ _ _ _ _ _ _ 1 _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
_ _ _ _ _ _ _ _ _ _ 1 _ _ _ _ _ _
_ _ _ 2 _ _ _ _ _ _ 2 _ _ _ _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 4
_ 58 7 476 56 19 6 3 5 6 12 6 2 1 2 41 12
_ _ _ _ _ _ _ _ _ _ _ _ _ 1 _ _ _
1 4 _ 9 1 2 _ _ _ _ _ _ 3 3 _ 3 _
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 2
_ _ _ _ _ _ _ _ _ _ 2 4 _ _ _ _ _
108 S.A. Kuzmina et al.
sample eco
Ikpikpuk B-0- Titaluk B-0-
taxon code
1 2 4 4a 5 1 2 3 4
Curimopsis sp. dt _ _ _ _ _ _ _ _ _
Fam. Coccinellidae
Subfam. Coccinellinae
Tribe Cephaloscymnini
Ceratomegilla ulkei Crotch, 1873 oth _ _ _ _ _ _ _ _ _
Tribe Coccinellini
Coccinella fulgida Wats., 1954 oth _ _ _ _ _ _ _ _ 1
C. transversoguttata Fald., 1835 oth _ _ _ _ 1 _ _ 1 _
Coccinella sp. oth _ _ _ 1 _ _ _ _ _
Fam. Chrysomelidae
Subfam. Chrysomelinae
Chrysolina (Arctolina) magniceps
(Sahlb., 1887)? ar
_ _ _ _ _ _ _ _ _
Ch. (Arctolina) septentrionalis
(Men, 1851) mt _ _ 1 3 5 6 2 1 _
Ch. (Arctolina) subsulcata (Mnnh.,
1853) ar _ _ _ _ 2 _ _ _ _
Ch. (Chalcoidea) marginata (L,
1758) mt? _ _ _ _ _ _ _ _ _
Ch. (Pleurosticha) tolli Jac., 1990 ar _ _ _ _ _ 3 5 2 _
Ch. (Allohypericia) basilaris (Say,
1824) mt? _ _ _ 1 _ _ _ _ _
Chrysolina sp. mt? _ _ _ _ _ _ _ _ _
Chrysomela blaisdelli (V.D., 1938) sh _ _ _ _ _ _ _ _ _
Phratora kenaiensis Brown, 1952 sh _ _ _ _ _ _ _ _ _
Phaedon armoraciae (L., 1758)? mt _ _ _ _ _ _ _ _ _
Subfam. Galerucinae
Tribe Galerucini
Galerucella stefanssoni (Brown,
1938) mt _ _ _ _ 2 _ _ _ _
Fam. Brentidae
Subfam. Apioninae
Mesotrichapion alaskanum (Fall,
1926) dt _ _ 1 _ _ _ _ 1 _
M. cyanitinctum (Fall, 1926) dt 1 _ 1 9 1 2 1 4 _
Apioninae gen. indet. dt? _ _ _ _ _ _ _ 1 _
Fam. Brachyceridae
Subfam. Erirhininae
Erirhinus aethiops (F., 1792) mt _ _ _ _ 2 _ _ _ _
Fam. Curculionidae
Subfam. Ceutorhynchinae
Ceutorhynchus subpubescens LeC.,
1876 st _ _ _ _ _ _ _ _ _
109Late Quaternary insects and freshwater invertebrates of the Alaskan North Slope
Table 2 (continued).
Таблица 2 (продолжение).
5 6 7 7a 8a 8b 9 11 11a 12 13 13a 14 15 16 17 18
_ _ _ _ _ _ _ 1 1 _ _ _ _ _ _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 1 _
_ _ _ _ _ _ _ _ _ 1 _ _ _ _ _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ 1 _ _ _ _
_ _ _ _ _ _ _ _ 1 _ 1 1 _ _ _ _ _
_ _ _ _ _ _ _ _ _ _ 1 _ _ _ _ _ _
_ 6 2 112 5 3 1 _ 1 2 16 10 16 15 7 2 _
_ _ _ 7 _ _ _ _ _ 1 _ _ _ 10 7 _ _
_ _ _ 10 1 _ _ _ _ _ _ 7 1 14 3 3 2
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
_ 2 _ 20 1 _ _ _ 1 _ 3 1 _ 1 2 2 _
_ _ _ _ _ _ _ 1 _ _ _ _ _ _ _ _ _
_ _ _ 2 _ _ _ _ _ 6 _ 2 _ _ _ 1 _
_ _ _ _ _ _ _ _ _ _ 1 _ _ _ _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ 1 _ _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
_ 4 _ 10 3 2 _ _ 1 _ 5 3 1 4 _ 1 _
_ 8 2 6 1 2 1 _ 1 _ _ 7 2 2 _ 4 2
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
_ _ _ 1 _ _ _ _ _ _ 1 _ _ _ 3 _ _
110 S.A. Kuzmina et al.
sample eco
Ikpikpuk B-0- Titaluk B-0-
taxon code
1 2 4 4a 5 1 2 3 4
Subfam. Entiminae
Tribe Alophinini
Lepidophorus lineaticollis Kby.,
1837 dt _ _ 8 16 93 3 2 12 2
L. thulius (Kiss., 1974) dt _ _ _ 1 3 _ _ _ _
Tribe Sitonini
Sitona aquilonius Bright, 1994 dt _ _ _ 1 7 _ _ _ _
S. lineellus (Bonsd., 1785) dt _ _ _ _ 1 _ _ _ _
Subfam. Hyperinae
Hypera castor (LeC., 1876) dt _ _ _ _ _ _ _ _ _
H. diversipunctata Schrank, 1798 dt _ _ 1 _ 2 _ _ _ _
H. trivittata (Say., 1831) dt _ _ _ 2 1 _ _ _ _
Hypera sp. dt _ _ _ _ _ _ _ _ _
Subfam. Lixinae
Tribe Cleonini
Stephanocleonus confusus (And.,
1987) st _ _ _ _ _ _ _ _ _
Coniocleonus parshus (And., 1987) st _ _ _ _ _ _ _ _ _
C. zherichini T.-M. et Kor., 1977 dt _ _ 3 _ 9 _ _ 4 _
Coniocleonus sp. st? _ _ _ _ _ _ _ _ _
Subfam. Molytinae
Tribe Hylobiini
Hylobius pinicola (Coup., 1864 ) _ _ _ _ _ _ _ 1 _
Tribe Lepyrini
Lepyrus canadensis Csy., 1895 sh _ _ _ _ 4 _ _ _ _
L. gemellus Kby., 1837 sh _ _ 1 _ 3 _ _ _ _
L. nordenskioeldi Faust, 1885 sh _ _ 2 2 22 1 _ 2 _
L. labradorensis Blair, 1933? sh _ _ _ _ _ _ _ _ _
Lepyrus spp. sh _ _ _ _ _ _ _ _ _
Subfam. Curculioninae
Tribe Rhamphini
Isochnus arcticus Kor., 1977 ar 2 _ 1 6 1 7 2 12 1
Subfam. Scolytinae
Tribe Halastini
Scierus annectans LeC., 1876 fo _ _ _ _ _ _ _ _ _
Tribe Scolytini
Scolytus piceae (Swaine, 1910) fo _ _ _ _ _ 1 _ _ _
Ord. Heteroptera
Fam. Corixidae
Sigara sp. r&a _ _ _ _ _ 1 _ _ _
111Late Quaternary insects and freshwater invertebrates of the Alaskan North Slope
Table 2 (continued).
Таблица 2 (продолжение).
5 6 7 7a 8a 8b 9 11 11a 12 13 13a 14 15 16 17 18
1 26 6 699 33 12 7 4 4 6 26 40 7 85 18 47 65
_ _ 1 8 3 2 _ _ _ _ _ _ _ 2 _ 1 1
_ 2 _ 9 2 2 _ _ _ 1 4 3 _ 1 _ 1 4
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 1 _
_ _ _ _ _ _ _ _ _ _ 1 _ _ _ _ _ _
_ 1 _ 2 1 _ _ _ _ _ _ 1 _ 1 _ 1 _
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
_ _ _ _ _ 1 _ _ _ _ _ _ _ _ _ _ _
_ _ _ _ 5 1 _ _ _ _ 2 3 _ 3 _ _ _
_ _ _ 11 _ _ _ _ _ _ _ _ _ 1 _ _ 16
_ 2 1 8 _ _ _ _ 1 2 _ _ 1 _ 3 4 7
_ _ _ 49 _ _ _ _ _ _ _ _ _ _ _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ _ 1 _ _ _
_ _ _ 12 1 _ _ _ _ _ 2 _ _ _ _ _ _
_ _ _ 6 _ _ _ _ _ 1 2 _ _ 3 2 _ _
_ 5 _ 69 3 _ _ 1 3 3 11 _ 4 7 4 4 25
_ _ _ _ _ _ _ _ _ 1 _ _ _ _ _ _
_ _ 1 _ _ 1 1 _ _ _ _ _ _ _ _ _ _
_ 12 3 12 1 7 _ _ 1 1 11 9 8 25 _ 3 1
_ _ _ _ _ _ _ _ _ _ _ _ _ 1 _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
_ 1 _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
112 S.A. Kuzmina et al.
sample eco
Ikpikpuk B-0- Titaluk B-0-
taxon code
1 2 4 4a 5 1 2 3 4
Fam. Saldidae
Saldula sp. r&a _ _ _ _ _ _ _ _ _
Chiloxanthus sp. r&a _ _ _ _ _ _ _ _ _
Ord. Trichoptera
Trichoptera larvae r&a _ _ _ _ _ _ _ _ _
sum
4 3 139 246 371 218 98 201 27
nus is regarded as valid again (Meregalli, Fre-
muth, 2013; Meregalli, 2017) but confusion
concerning North American species still exists.
Recently (Meregalli, 2017) part of the North
American Stephanocleonus has been returned
to Coniocleonus: Stephanocleonus stenotho-
rax And., 1987 = Coniocleonus zherichini T.-
M. et Kor., 1977; Stephanocleonus parshus
And., 1987 = Coniocleonus parshus (And.,
1987); Stephanocleonus plumbeus LeC., 1876
= Coniocleonus plumbeus (LeC). Two North
American Stephanocleonus: S. confusus And.,
1987 and S. cristatus LeC., 1876 remained in
this genus.
2. dt — insects of dry, open habitats in the
tundra and partly in boreal forest: Lepidophorus
lineaticollis, Coniocleonus zherichini, Sitona
aquilonius Bright, 1994, Hypera diversipunc-
tata (Schrank, 1798), Amara alpina, Stereocer-
us haematopus and others.
3. mt — insects associated with mesic to
moist tundra habitats or meadows and bogs in
boreal forest. These include Pterostichus (Cry-
obius) spp., Tachinus brevipennis Sahlb., 1880,
Chrysolina septentrionalis (Men., 1851), and
others.
4. ar — Arctic insects. There are cold-resis-
tant beetles, living today in the northern tundra
and polar desert: Chrysolina subsulcata (Mann.,
1853), Ch. tolli (Jac., 1910), Isochnus arcticus
Kor., 1977.
5. sh — insects that live and feed on shrubs,
mostly willows: Lepyrus spp., Chrysomela spp.
and others.
6. pl — plant litter inhabitants including
many species of rove beetles (Lathrobium spp.,
Quedius spp.) and Leiodidae
7. fo — forest species. Here the group is
represented by two species of bark beetles (Sci-
erus annectans LeC., 1876, Scolytus piceae
(Swaine, 1910)).
8. r&a — riparian and aquatic insects
(Elaphrus spp., Bembidion umiatense Lindr.,
1963, Agabus moestus (Curt., 1835)
9. oth — other insects, without certain eco-
logical status, such as lady birds Coccinella spp.
We calculated the minimum number of indi-
viduals (MNI), based on the maximum number
of any individual sclerite (head, pronotum,
elytron) of a taxon in a sample. All MNI of
certain ecological group are summed up. The
result is graphic of ecological change of the
insect assemblages thought the section (Fig. 6).
Results
Fossil insects from the area have previously
been studied by R. Nelson. He made his Ph.D.
dissertation (Nelson, 1982, 1986) on a study of
Pleistocene and Holocene fossil insects from
sites on the Ikpikpuk and Titaluk Rivers. He
presented his results in meetings (Nelson, Cart-
er, 1997; Nelson, 1998) and published a de-
tailed paper about the insects of the Populus
Beds from the Ikpikpuk section called Cotton-
wood bend (Nelson, Carter, 1987). Until now,
no further work on the fossil insects of the North
Slope has been done, despite their abundance
and good preservation.
113Late Quaternary insects and freshwater invertebrates of the Alaskan North Slope
Table 2 (continued).
Таблица 2 (продолжение).
5 6 7 7a 8a 8b 9 11 11a 12 13 13a 14 15 16 17 18
_ _ 2 _ _ 1 _ _ _ _ _ _ _ _ _ _ _
_ _ _ _ _ _ _ _ _ 1 _ _ _ 2 _ _ 1
1 _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
4 346 92 3114 272 165 26 34 43 105 397 348 506 639 186 254 207
This beetle is common in modern tundra, relict
steppe and openings in the boreal forest in
northern North America. Recently the species
has been recorded from Chukotka (Berman et
al., 2002); in the Pleistocene it lived only in
Eastern Beringia (Kuzmina, Matthews, 2012).
L. lineaticollis indicates dry tundra or steppe
tundra environments (Matthews, 1983).
Xerophilous insects dominate only one as-
semblage, IK-B05 (49%) in the lower unit;
other assemblages yield up to 52% mesic tundra
species, with the most numerous being speci-
mens of Pterostichus (Cryobius) and the rove
beetle Tachinus brevipennis. However, all the
samples except T-B01 and T-B02 contain spe-
cies associated with of steppe-tundra environ-
ments, especially Morychus aff. aeneolus. All
assemblages yielded members of the arctic
group, including the weevil Isochnus arcticus
and the leaf beetle Chrysolina tolli. Curiously,
sample T-B01 yielded a clear indicator of conif-
erous forest, the spruce bark beetle Scolytus
piceae. Such a strange combination of arctic
and forest beetles could have been caused by
rapid changes of environment and accumulation
of the beetle remains in an alluvial deposit. Part
of the fossil assemblage was probably reworked
but in any case the presence of forest beetle in
this region suggests at least a short warming
interval.
Quaternary deposits in the Ikpikpuk-Titaluk
area contain tree remains; there are big willow
stumps in growth position, as well as poplar
stumps and leaves. Poplar remains were found
mostly in Populus Beds, but several willow
The fossil insects from Titaluk and Ikpikpuk
are presented in Table 2 and Figs 7, 8.
Insects from the lower gravel (A) come from
both sites (Figs 3–6). Samples Ikpikpuk 1 and 2
are poor, sample 3 is empty, but samples 4, 4a
and 5 yielded good results. The lower unit was
sampled from the banks of the Titaluk River in
3 profiles; all 5 samples are rich (up to 500
MNI).
Tundra species are represented very well:
there are several species of the typical tundra
ground beetle subgenus Pterostichus (Cryo-
bius), including P. brevicornis, P. ventricosus,
P. pinguedineus, P. caribou, P. nivalis, P. tareu-
miut, and others. Abundance and diversity of
Cryobius is observed in each studied samples.
Remains of other cold-adapted ground bee-
tles include Amara (Curtonotus) alpina, Pteros-
tichus (Tundraphilus) sublaevis, and Stereo-
cerus haematopus. These taxa are abundant
throughout the whole section.
The relict tundra ground beetle Poecilus
(Derus) nearcticus was found here as well as in
other North Slope assemblages. This species is
common in fossil assemblages in both Eastern
and Western Beringia. The species was origi-
nally described (Lindroth, 1966) from the Ander-
son River Delta. Later it was discovered in
northeastern Siberia (Kiselev, 1981). We found
modern specimens on a sandy slope with scarce
vegetation at the Titaluk River. The modern
distribution of P. nearcticus reflects its Bering-
ian origin.
The most common beetle in the whole sec-
tion is the weevil Lepidophorus lineaticollis.
114 S.A. Kuzmina et al.
Fig. 7. Fossil insects (Carabidae — Coccinellidae) from the Titaluk river site.
1 — Carabus truncaticollis, 2 — Elaphrus parviceps, 3 — Bembidion arcticum, 4 — B. sordidum, 5, 6 — Pterostichus
arcticola, 7, 8 — P. brevicornis, 9, 10 — P. empetricola 11 — P. pinguedineus, 12 — P. tareumiut, 13, 14 — P.
ventricosus, 15 — P. sublaevis, 16, 17 — Stereocerus haematopus, 18, 19 — Amara alpina, 20 — Cymindis unicolor,
21 — Hydroporus lapponum, 22 — H. polaris, 23 — Oreodytes laevis, 24 — Agabus confinis, 25 — A. serricornis;
26 — Colymbetes dolabratus, 27 — C. dahuricus, 28 — Leiodes collaris, 29 — Agathidium angulare, 30 —
Micralymma brevilingue, 31 — Olophrum rotundicolle, 32 — Tachinus brevipennis, 33 — Tachinus sp., 34 — Stenus
sp., 35, 36 — Philonthus subvirescens, 37 — Lathrobium sibiricum, 38 — Quedius sublimbatus, 39 — Helophorus
splendidus, 40, 41 — Cymbiodyta sp., 42, 43, 44 — Aphodius pectoralis, 45, 46 — Curimopsis albonotata, 47 —
Ceratomegilla ulkei, 48 — Coccinella transversoguttata; 1 — T03–B17; 2, 3, 4, 5, 6, 7, 8, 11, 12, 13, 14, 17, 18, 19,
24, 26, 35, 36, 37 — T03–B13; 9, 10, 21, 22 — T03–B07a; 16, 29, 31, 32, 33, 40, 41 — T03–B15; 20, 34 — T03–B14;
23, 25 — T03–B12; 27 — T03–B11a; 28, 45, 46 — T03–B06; 30 — T03–B11; 38 — T03–B04; 39, 48 — T03–B03;
115Late Quaternary insects and freshwater invertebrates of the Alaskan North Slope
42, 43, 44, 47 — T03–B17; 1, 2, 4, 21, 22, 24, 25, 27, 28, 38, 39, 40, 44, 46, 47 — right elytron; 3, 15, 17, 19, 23, 29,
33 — left elytron; 6, 8, 10, 14 — both elytra; 5, 7, 9, 11, 12, 13, 16, 18, 20, 26, 30, 31, 34, 37, 40, 43, 45, 48 — pronotum;
35, 42 — head. Scale bar 1 mm.
Рис. 7. Ископаемые насекомые (Carabidae — Coccinellidae) из разреза на реке Титалук.
1 — T03–B17; 2, 3, 4, 5, 6, 7, 8, 11, 12, 13, 14, 17, 18, 19, 24, 26, 35, 36, 37 — T03–B13; 9, 10, 21, 22 — T03–B07a;
16, 29, 31, 32, 33, 40, 41 — T03–B15; 20, 34 — T03–B14; 23, 25 — T03–B12; 27 — T03–B11a; 28, 45, 46 — T03–
B06; 30 — T03–B11; 38 — T03–B04; 39, 48 — T03–B03; 42, 43, 44, 47 — T03–B17; 1, 2, 4, 21, 22, 24, 25, 27, 28,
38, 39, 40, 44, 46, 47 — правое надкрылье; 3, 15, 17, 19, 23, 29, 33 — левое надкрылье; 6, 8, 10, 14 — соединенные
надкрылья; 5, 7, 9, 11, 12, 13, 16, 18, 20, 26, 30, 31, 34, 37, 40, 43, 45, 48 — переднеспинка; 35, 42 — голова. Шкала
1 мм.
stumps and a single poplar stump were found in
the lower gravel (Mann et al., 2010). Spruce
remains are rare there. A few fragmental spruce
needles were observed in the Populus Beds
(Nelson, Carter, 1987); they are poorly pre-
served, highly oxidized and are believed to be
reworked. Our team found a spruce stump on a
sandy bar on the Titaluk River. So, spruce grew
in the area in some period of the Pleistocene, but
this was an exceptional episode.
Reworking of fossil insects likely took place
here. Usually erosion, flotation, coarse gravel
and other impacts destroy fragile chitin frag-
ments. But here we can see some evidence of
possible reworking. Coarse alluvium of the low-
er unit contains well- preserved chitin and the
concentration of beetle fragments is extremely
high. These fragments were probably sorted by
size, for example big and heavy elytra of Le-
pyrus were concentrated in a cross-bedded layer
with coarse gravel (sample T-B13) while they
are absent in the sample T-B13a which was
taken 10cm higher, from sediments containing
tiny plant debris. Local chitin preserved under
permafrost conditions is not fragile.
Perhaps the repetitive ecological composi-
tion of the insect assemblages throughout the
whole section (Fig. 6, Table 2) could be partly
explained by reworking.
The proportion of riparian and aquatic in-
sects is low in the gravel unit. Aquatic species
are represented by the widespread arctic diving
beetles Agabus moestus, Colymbetes dolabra-
tus, various species of Hydroporus, and the
water scavenger beetle Helophorus splendidus.
These beetles prefer small vegetated standing
water, not a fast flowing river. Probably they
occupied pools in the river floodplain. The
floodplain also provided habitats for Bembidi-
on umiatense, Olophrum latum, O. rotundicol-
le, Stenus sp. and others.
The main sandy unit (B) was sampled in 4
profiles of the Titaluk section (Figs 4–6). The
abundance of insect fossils varied greatly be-
tween samples (Table 2) from almost none to
very rich. The richest sample (T-B07a) con-
tained a minimum of 3114 individuals. The
layer from which the samples were taken was
full of insect remains; it was a very rare case in
which insect chitin actually formed the bulk of
the sediment.
Insect assemblages taken from the sand unit
are more-or-less similar to those described
above. However, there is evidence of increasing
environmental aridity with age, because the
proportion of dry tundra and steppe-tundra
groups became more prominent in the upper
samples. Weevils play a more significant role
than before, especially xerophilous species such
as Sitona aquilonius, Hypera diversipunctata,
Stephanocleonus confusus, Coniocleonus
parshus, and C. zherichini. These species are
quite rare in the modern environment but were
common in the Pleistocene. The most interest-
ing discovery was the finding of S. confusus and
C. parshus. The first species is known today
from southern Alberta and Saskatchewan and
south to Arizona; the second is widely distribut-
ed in North America as far north as northern
Alberta and British Columbia, but does not
reach Alaska. Neither species has been record-
ed from Arctic tundra. The third species, C.
zherichini, has a trans-Beringian distribution
today, but this species is quite rare in modern
collections. These southern weevils, in combi-
nation with extremely abundant fossils of Lep-
idophorus lineaticollis, the exotic Morychus
aff. aeneolus, and the relict Poecilus nearcti-
116 S.A. Kuzmina et al.
Fig. 8. Fossil insects (Chrysomelidae — Saldidae) from the Titaluk river site.
1 — Chrysolina basilaris, 2 — C. septentrionalis, 3 — C. magniceps, 4 — C. subsulcata, 5 — Chrysolina sp., 6 —
Chrysomela blaisdelli, 7 — Phratora kenaiensis, 8 — Mesotrichapion alaskanum, 9 — M. cyanitinctum, 10, 11, 12 —
Sitona aquilonius, 13, 14 — Lepidophorus thulius, 15 — Hypera castor, 16 — H. diversipunctata, 17, 18 —
Stephanocleonus confusus, 19, 20 — Coniocleonus parshus, 21, 22 — C. zherichini, 23 — Coniocleonus sp., 24 —
Lepyrus nordenskioeldi, 25 — Isochnus arcticus, 26 — Ceutorhynchus subpubescens, 27 — Chiloxanthus sp.; 1 —
T03–B06; 2, 3, 7, 15, 16, 17, 18, 24, — T03–B13; 4 — T03–B15; 8, 9 — T03–B13a; 10, 11, 13, 19, 20, 21, 22, 23 —
T03–B18; 12, 14, 26 — T03–B07a; 25 –T03–B03; 27 — T03–B12; 1, 3, 6, 8 — right elytron; 2, 7, 12, 15, 16, 25, 27 —
left elytron; 4, 14 — both elytra; 4, 9, 13, 23 — pronotum; 10, 11, 17, 18, 19, 20, 21, 22, 24 — head. Scale bar 1 mm.
117Late Quaternary insects and freshwater invertebrates of the Alaskan North Slope
Рис. 8. Ископаемые насекомые (Chrysomelidae — Saldidae) из разреза на реке Титалук.
1 — T03–B06; 2, 3, 7, 15, 16, 17, 18, 24, — T03–B13; 4 — T03–B15; 8, 9 — T03–B13a; 10, 11, 13, 19, 20, 21, 22,
23 — T03–B18; 12, 14, 26 — T03–B07a; 25 –T03–B03; 27 — T03–B12; 1, 3, 6, 8 — правое надкрылье; 2, 7, 12,
15, 16, 25, 27 — левое надкрылье; 4, 14 — соединенные надкрылья; 4, 9, 13, 23 — переднеспинка; 10, 11, 17, 18,
19, 20, 21, 22, 24 — голова. Шкала 1 мм.
cus, make the fossil Pleistocene assemblages
quite peculiar. There is certainly no modern
analogue for this combination of species.
Single specimens of forest beetles were found
in the lower part of the sandy unit. The sample
T-B15 contains the single remains of the weevil
Hylobius pinicola and the bark beetle Scierus
annectans, and T-B03 also contained H. pinico-
la. These species are indicators of coniferous
forest. Despite the possibility of reworking, we
are forced to conclude that a warming episode
took place in the region sometime during the
Pleistocene.
The Yedoma-analogue unit (C), in contrast
with lower gravel and sand, seemed devoid of
insect remains, but by screening a couple of
hundred kg of sediment we obtained two good
assemblages from three samples (Figs 5–6, Ta-
ble 2). The proportion of the steppe-tundra
group in sample T-B08a is highest in the section
— 24%. Otherwise, the species composition
and general ecological structure of these two
assemblages was similar to the previous ones.
The Poplar Beds unit (D) was found in two
profiles at Titaluk River (Figs 4–6, Table 2). In
spite of the climatic amelioration that apparent-
ly allowed poplar to invade the North Slope, the
general species and ecological composition of
the insect assemblages remained the same. A
similar picture has been observed in the insect
fossil assemblages from the Populus Beds of the
Ikpikpuk River (Nelson, Carter, 1987). Xero-
philous insects dominated most of the assem-
blages, and the steppe-tundra still group played
significant role. The Arctic group became less
evident but single remains of Isochnus arcticus
were still present in each sample. The main
difference was observed in samples T-B10, 11,
and 12 from the first profile (Fig. 4). This
section consisted of bluish-gray fine sand with
layers of well-preserved poplar leaves. We not-
ed a sharp increase in the water-riparian group,
especially in sample T-B11 (Fig. 6, Table 2).
They are represented mostly by the diving bee-
tles of the genera Agabus and Hydroporus.
Sample T-B12 is only one in the section that
contained fossils of the winter eggs (ephippia)
of water fleas (Fig. 9). We recognized numerous
ephippia of Daphnia cf. longispina O.F. Müller
and Simocephalus sp. (Fam. Daphniidae Straus).
Exact species identification of ephippia is diffi-
cult (Kotov, 2015). Different taxa of the D.
longispina can live in a variety of fresh water
habitats (Benzie, 2005). Members of the genus
Simocephalus are typical inhabitants of shal-
low, vegetated margins of standing water (Orlo-
va-Bienkowskaja, 2001). The fossils water fleas
indicate the presence of temporary floodplain
pools oxbows, but not the river itself.
The second profile (Fig. 5) probably repre-
sents a river channel. It overlay the older unit
with a clear unconformity. Dynamic fluvial con-
ditions supported reworking and mixing of old-
er and younger fossils.
Discussion
Nelson (1982, 1986) started the investiga-
tion of North Slope fossil insects and vegetation
in the 1980s. He proposed the following scenar-
io of environmental changes:
1 — before 40 ka: Environment similar to
modern with less abundant shrub vegetation and
more open surfaces. Climate was cooler and
drier than today.
2 — between 40 and 25 ka: decreasing
moisture and temperature.
3 — Early Holocene. Populus balsamifera
and a few other species were present north of
their modern limits. Climate was warmer than
today but the landscape was discontinuously
vegetated and summer precipitation may have
been infrequent.
Our climate estimation using MCR method
is shown in the Table 3. In general, the recon-
118 S.A. Kuzmina et al.
Fig. 9. SEM photos of daphniid ephippia (Cladocera) from sample T03-B-12.
A–B — Daphnia cf. longispina C–D — Simocephalus sp.; A, C — general view, B, D — surface sculpture.
Fig. 9. Фотографии со сканирующего электронного микроскопа эфиппиумов дафнид (Cladocera) из
образца T03-B-12.
A–B — Daphnia cf. longispina; A, C — общий вид, B, D — скульптура поверхности.
struction shows lower than recent temperatures
during all time intervals including the Early
Holocene. The warming signal in the sample T-
B-09 is not much reliable because the examined
insect assemblage is small. Another assemblage
what provides warmer than recent temperature
T-B-14 from the lower unit A is big but thermo-
philous species which affected the MCR result
is only one — Cymindis unicolor. This species
is more likely absent in the studied area recently,
R Nelson (Nelson, 2001) who worked exactly at
the Titaluk River as well as in other sites of the
North Slope did not collected it; we also did not
find the species in the modern environment. C.
unicolor lives in the tundra and alpine treeless
environment on well-drained soil. It occurs from
Newfoundland to southwest Yukon and in moun-
tains across Canada and western USA (Bousquet,
2012).
The rest of species composition is similar to
these of other samples. From the other hand we
found several forest species in the lower unit.
These single evidences indicate probably erod-
ed interglacial layer.
Climate estimation of the Poplar Beds using
fossil insects shows lower temperature (7.5 to
10°C than mean July temperatures 14 to 15°C
reconstructed using Populus balsamifera (Nel-
son, Carter, 1987). Thermophilous insects did
not follow the poplar groves, and it is hard to
explain. Maybe the Brook Ridge prevents the
spreading of southern fauna to this area while
poplar seeds which are adapted to wind trans-
portation were able to overcome this barrier.
119Late Quaternary insects and freshwater invertebrates of the Alaskan North Slope
Table 3. Reconstruction using MCR of mean July (TMAX), mean January (TMIN), and differences with
modern temperature of the nearest weather station — Umiat, AK (69.22°N, 152.08°W).
Таблица 3. Реконструкция методом MCR среднеиюльских температур (TMAX) и среднеянварских
(TMIN) и разница с современной температурой по данным ближайшей метеостанции Умиат,
Аляска (69.22°N, 152.08°W).
recent 10 –30
Sample TMAX TMIN
No. spp
used TMAX TMIN
Ikpikpuk B-0-4 Unit A 7.5–9.5 –33.5 to –28.5 15 –2.5 to –0.5 –3.5 to +1.5
Ikpikpuk B-0-4a Unit A 8.5–9 –31 to –29.5 20 –1.5 to –1 –1 to +0.5
Ikpikpuk B-0-5 Unit A 8–9.5 –33 to –28.5 14 –2 to –0.5 –3 to +0.5
Titaluk B-0-1 Unit A 5.5–8 –31.5 to –26 16 –4.5 to –2 –1.5 to +4
Titaluk B-0-2 Unit A 7–9 –31.5 to –24 10 –3 to –1 –1.5 to +6
Titaluk B-0-13 Unit A 9–10 –31.5 to –28.5 22 –1 to 0 –1.5 to +1.5
Titaluk B-0-13a Unit A 8.5–9 –31 to –29 21 –1.5 to –1 –1 to +1
Titaluk B-0-14 Unit A 11–11.5 –21.5 to –20.5 15 +1 to +1.5 +8.5 to +9.5
Titaluk B-0-3 Unit B 8.5–9 –31 to –29.5 17 –1.5 to –1 –1 to +0.5
Titaluk B-0-4 Unit B 8–10 –31 to –22 9 –2 to 0 –1 to +8
Titaluk B-0-6 Unit B 8.5–9 –31 to –29.5 15 –1.5 to –1 –1 to +0.5
Titaluk B-0-7 Unit B 8.5–9 –31 to –29.5 12 –1.5 to –1 –1 to +0.5
Titaluk B-0-7a Unit B 8.5–9 –31.5 to –29 24 –1.5 to –1 –1.5 to +1
Titaluk B-0-15 Unit B 8–9 –31 to –29 28 –2 to –1 –1 to +1
Titaluk B-0-8a Unit C 8–8.5 –32 to –30.5 18 –2 to –1.5 –2 to –0.5
Titaluk B-0-8b Unit C 8.5–9 –31 to –29.5 18 –1.5 to –1 –1 to –0.5
Titaluk B-0-9 Unit C 9.5–10 –25 to –22 6 –0.5 to 0 +5 to +8
Titaluk B-0-11 Unit D 7.5–9 –32 to –29 9 –2.5 to –1 –2 to +1
Titaluk B-0-11a Unit D 8.5–9 –31 to –29 8 –1.5 to –1 –1 to +1
Titaluk B-0-12 Unit D 8.5–9 –31 to 29.5 13 –1.5 to –1 –1 to +0.5
Titaluk B-0-16 Unit D 8–10 –33 to –25 10 –2 to 0 –3 to +5
Titaluk B-0-17 Unit D 8.5–9.5 –31 to –26.5 17 –1.5 to –0.5 –1 to +3.5
Titaluk B-0-18 Unit D 9–9.5 –30 to –29 11 –1 to –0.5 0 to +1
Our results support Nelson’s conclusions
about drier-than-modern climate and warming
in the early Holocene. He noticed that the main
surface (away from the river valley) was proba-
bly unstable, dry, sandy, and covered by discon-
tinuous vegetation. Such surface conditions also
existed during cold Pleistocene intervals, as
well as in the early Holocene (Nelson, Carter,
1987).
The difference between our reconstructions
and those of Nelson is that he never used the
term “steppe-tundra”. This concept became
much more widely discussed in the 1990s, well
after Nelson’s dissertation was finished. Per-
haps he was also influenced by the plant macro-
fossil and pollen results, as plant remains rarely
indicate the steppe-tundra environment. Also,
his list of identified insects only included tundra
species, and Coniocleonus specimens were rare
in his assemblages, and were simply identified
as Cleonus sp. The taxonomic understanding of
this group of weevils has advanced consider-
ably in the last 40 years. Our specific identifica-
tion of Stephanocleonus confusus and Conio-
120 S.A. Kuzmina et al.
cleonus parshus shows that some species were
present north of their modern limits, even dur-
ing the Pleistocene. This situation is more typ-
ical of Western Beringian steppe-tundra ento-
mofaunas (Kiselev, 1981; Kuzmina, 2015), of
which Nelson had little or no knowledge during
the 1980s, when nearly all Russian research in
this field was published in the Russian language,
and was virtually unobtainable in the US.
The abundance of xerophilous species, es-
pecially Morychus and Lepidophorus lineati-
collis has been recorded in the Ikpikpuk (Nel-
son, Carter, 1987) and Titaluk Populus Beds.
The Titaluk faunal succession suggests that the
insect fauna changed little at the Pleistocene-
Holocene transition. Steppe-tundra environment
survived the early Holocene warming in this
region, perhaps because of the cold climate and
lack of well developed forest in this northern
locality. A similar situation was observed in
northern localities in Western Beringia (An-
dreev et al., 2009; Kuzmina, 2015) while in
southern regions, such as Klondike Gold Field,
the Pleistocene-Holocene transition is sharp and
the steppe-tundra fauna disappeared from the
record almost completely (Kuzmina, unpub-
lished).
Single forest insects, wood and stumps were
found occasionally in the Pleistocene part of the
section. We can not correlate them with specific
layers, and perhaps most of them were reworked
from older deposits of unknown age. It is prob-
able that forest-associated insect remains were
eroded out of sediments from a Pleistocene
warm interval, such as the Last Interglacial. It is
hard to verify easily because the radiocarbon
dates of the wood from the lower units (Table 1)
are infinitive or close to the method limit.
Open, sandy surfaces helped to maintain
high soil temperatures that are critical for the
development of many thermophilous insects.
For instance Cleonini weevils are very sensitive
to soil temperature (Berman et al., 2011). They
need significant amounts of warmth for the
development of larvae in the soil. Arctic regions
could provide the necessary sum of positive soil
temperatures during the summer months when
daylight hours are long or continuous. The ab-
sence of trees and moss cover is also critical for
the maintenance of soil warming. Under a dry
Pleistocene – Early Holocene climate regime,
thermophilous Cleonini weevils flourished in
the northern Beringian steppe-tundra.
Continuous eolian sand deposition played
an important role in the local environment of the
study sites. Late Pleistocene eolian activity af-
fected the region widely. As discussed above, a
large sand dune area (sand sea) is located very
exactly near to the studied sections (Carter,
1981). Another sand deposit of eolian origin,
the Kittigazuit Formation, is known from the
Tuktoyaktuk Coastlands in the northern part of
the Canadian Northwest Territories (Murton,
1996). Fossil insect assemblages from Kittiga-
zuit Formation (Dallimore et al., 1987, Murton
et al., 2017) are similar in species composition
to those found in the Ikpikpuk-Titaluk area.
Wind itself helped to maintain favourable
environmental conditions for key Pleistocene
insect species. Observations of Berman (1990)
in northeastern Siberia allow us to reconstruct
the environmental requirements of the key Pleis-
tocene species Morychus viridis Kuzm. et Ko-
rot., 1987. This pill beetle feeds on certain
minute mosses that can develop only on dry
open surfaces. Wind helps to develop such
surfaces by preventing the development of the
dense moss-lichen cover found in the tundra and
in the boreal forest. Eastern Beringian M. aff.
aeneolus probably occupied similar habitats to
M. viridis (Kuzmina, unpublished).
The long-term presence of such Arctic in-
sects as Isochnus arcticus and several species of
Chrysolina may seem to contradict the presence
of other faunal elements. This seeming contra-
diction may be explained in two different ways:
(1) Modern species with high Arctic distribu-
tion are relicts of the Pleistocene steppe-tundra
community. They, being cold-resistant, survive
now in northern open treeless landscapes where
they avoid competition with southern species.
Indeed, almost all Pleistocene steppe-tundra
assemblages contain Arctic species (Kuzmina,
Matthews, 2012). They were equitable mem-
bers of this non-analogue community. (2) Arctic
species were indifferent to climate but sensitive
121Late Quaternary insects and freshwater invertebrates of the Alaskan North Slope
for discontinuous vegetation cover. This sug-
gestion needs specification – the Arctic group is
present in most of the fossil assemblages, but it
only became dominant during the coldest Pleis-
tocene intervals (Sher et al., 2005) and even
then this dominance occurred mostly in north-
ernmost sites. Under this scenario, the arctic
species indicate substantial cooling if their num-
bers increase dramatically.
The task of “looking at the Eastern Bering-
ian section through Western Beringian eyes”
was successfully realized. We made certain that
the Siberian pill beetle, Morychus viridis, and
the North American fossil specimens of Mory-
chus are indeed different species. Furthermore,
we were able to verify that the North American
fossil Cleonini specimens belong to a regional
species, and that the diversity of Pterostichus
(Cryobius) species in Eastern Beringian fossil
insect faunas is greater than the diversity of this
group in Siberian faunas.
North Slope fossil insect assemblages show
some differences between Western and Eastern
Beringian steppe-tundra communities. Steppe
elements are more prominent in Siberia; the
Alaskan North Slope steppe-tundra was more
“tundra-like”. The dominant beetle taxon of the
local community is the weevil Lepidophorus
lineaticollis; Morychus plays significant but
secondary role. In Siberia L. lineaticollis is
absent, and Morychus viridis specimens often
comprise 80–90% of assemblage.
Many species recorded in the North Slope
fossil assemblages have Amphi-beringian dis-
tributions (i.e., they are found today on both
sides of the Bering Strait). These include Bem-
bidion umiatense, Dicheirotrichus mannerhei-
mi, Harpalus vittatus alaskensis, Poecilus ne-
arcticus, Pterostichus pinguedineus, P. brevi-
cornis, P. ventricosus, P. agonus, P. costatus,
P. vermiculosus, P. sublaevis, Stereocerus hae-
matopus, Amara alpina, Agabus moestus, Helo-
phorus splendidus, Micralymma brevilingue,
Tachinus brevipennis, Chrysolina septentrion-
alis, Ch. subsulcata, Hypera diversipunctata,
Coniocleonus zherichini, Lepyrus nordenskioel-
di, and Isochnus arcticus. Lepidophorus thulius
was Amphi-Beringian in the Pleistocene but
today it is found live only in North America.
Our study of Eastern Beringia fossil insects
has allowed us to recognize another important
feature. Many cold-phase Pleistocene insect
assemblages from the Yukon (Kuzmina, Mat-
thews, 2012; Matthews et al., 2019) are domi-
nated by the xerophilous weevil Connatichela
artemisiae And., 1984. This species has not
been found in any of the North Slope samples;
it is also absent in fossil assemblages from
Western Beringia. This weevil feeds on Artemi-
sia. It appears that Artemisia-dominated steppes
developed only in the interior regions of Eastern
Beringia during the Last glaciation.
The vegetation cover of the North Slope
steppe-tundra probably included various grass-
es, as indicated by the presence of Amara alpi-
na. The presence of herbs including legumes is
indicated by the presence of Coniocleonus, Si-
tona, and Hypera diversipunctata. Buttercups
are the host plant of Chrysolina septentrionalis,
and cruciferous plants are eaten by Chrysolina
tolli and Ceutorhynchus subpubescens. Tall
shrub willows are attacked by Lepyrus canaden-
sis, L. gemellus, L. nordenskioeldi, and Chry-
somela blaisdelli, while dwarf willow is the host
plant for Isochnus arcticus. The bark beetles
Hylobius pinicola, Scierus annectans, and Sco-
lytus piceae feed on coniferous trees.
Conclusions
This study is contributed to our knowledge
of the Pleistocene-Early Holocene insect faunas
of the Alaskan North Slope, and the fossil evi-
dence provided the basis for our paleoenviron-
mental reconstructions, as well as yielding new
information concerning the nature of the now-
extinct steppe-tundra community. The climate
during the Late Pleistocene and Early Holocene
allowed the steppe-tundra insect fauna to sur-
vive in situ, in spite of global warming at the end
of the Last Glaciation. Early Holocene warming
and the advance of poplar forest onto the North
Slope did not greatly affect the regional insect
fauna, perhaps because poplar trees were sparse
on the landscape, and did not shade all the
surfaces. Low moisture was probably more im-
122 S.A. Kuzmina et al.
portant in maintaining regional steppe-tundra
temperature regime.
Herbs and grasses dominated the local veg-
etation cover and Artemisia did not play a sig-
nificant role here. Tall willow shrubs occupied
riparian habitats, while dwarf willow probably
grew on upland surfaces. Wind-driven sand
deposition probably helped maintain the steppe-
tundra community on the North Slope.
The species and ecological requirements of
the North Slope fossil beetle assemblages indi-
cate a specific local environment that differed
from more southerly localities in Beringia, thus
suggesting regional zonation of Beringian eco-
systems.
The steppe-tundra insect fauna of the North
Slope is both similar to and different from those
described from Northeastern Siberia. The two
faunas share may species in common but the
xerophilous elements are mostly different. These
faunal differences were most likely due to dif-
ferences in the climates of the two regions. It
appears that northern Western Beringian cli-
mates were drier than those of northern Eastern
Beringia.
Acknowledgments
The field research was much assisted by the
National Parks Service of the USA. We thank
the Russian Foundation for Basic Research
(grants 04-04-48770 for SK), Leverhulme Foun-
dation, F/07 537T (grant for SE) and Russian
Government Program of Competitive Growth
of Kazan Federal University for support of
palaeoentomological studies. This study of Ber-
ingian cladocerans is supported by the Russian
Foundation for Basic Research (grant 18-04-
00398). We appreciate kindness of Pamela
Groves and Daniel Mann who recognized our
field location. Special thank to staff of the
Canadian national collection of insects, arach-
nids and nematodes (CNC) for their hospitality
and patience.
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Responsible editor K.G. Mikhailov
... Some body structures such as elytra or pronotum are not strong morphological features for Cryobius species taxonomy, and male genitalia are rarely found among fossil ground beetles. Fossil insects are being used as ecology indicators for Arctic paleoenvironmental reconstructions in the Pleistocene and the Holocene [3,[44][45][46][47][48]. Carabidae and particularly Cryobius frequently predominate in samples from different stratigraphic layers of Arctic territories, and they are associated with mesic to moist tundra habitats or meadows and bogs in boreal forest [47,48]. ...
... Some body structures such as elytra or pronotum are not strong morphological features for Cryobius species taxonomy, and male genitalia are rarely found among fossil ground beetles. Fossil insects are being used as ecology indicators for Arctic paleoenvironmental reconstructions in the Pleistocene and the Holocene [3,[44][45][46][47][48]. Carabidae and particularly Cryobius frequently predominate in samples from different stratigraphic layers of Arctic territories, and they are associated with mesic to moist tundra habitats or meadows and bogs in boreal forest [47,48]. For instance, in the taiga and tundra of North Europe, specimens of P. b. brevicornis have been sampled in broader environmental conditions [49][50][51][52][53]. Specimens of P. nivalis are usually found on loamy soil in rather dry tundra [19]. ...
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Subgenus Cryobius is one of the most numerous among the megafauna of tundra soils, but studies on its species distribution, taxonomy, and ecology are lacking. Phylogeny and phylogeography reconstructions of insects with taxonomic complexity have become possible using an integrative approach. Here, we report that specimens of Pterostichus (Cryobius) mandibularoides, described from North America, were detected in Eurasia. Thus, this species has a trans-Beringian range with high distributions in North America, as well as a disjunctive part of the range on the northeastern edge of Asia within Chukotka and Wrangel Island. Eight COI haplotypes with closed relationships (1–2 mutation steps) were detected within the whole range, and one 28S rRNA haplotype was detected for Eurasia. Bayesian phylogeny revealed that P. mandibularoides had the most recent common ancestor with sister species P. brevicornis and P. nivalis. Mean genetic distances of both. markers were similar and higher between P. mandibularoides and both P. brevicornis and P. nivalis (>5% ± 1.0%) than between the latter species (<4% ± 1.0%). The obtained results change the previous view about brevicornis group stock differentiation within Cryobius in the Arctic and require a revision of the phylogeny and phylogeography of brevicornis group species and Cryobius altogether.
... In some cases (e.g., Old Crow or Titaluksee Kuzmina et al., 2019) presence of ephippia in deposits in a significant number apparently is correlated with their lake-bog origin. Therefore, data on cladoceran analysis agree well with data from other types of the analysis. ...
... Этот выпуск журнала "Invertebrate zoology" посвящен его 80-летнему юбилею. Как цитировать эту статью: Kuzmina S.A., Elias S.A. 2019 area of his research was focused on the Kolyma Lowland. Andrei worked in collaboration with scientists from Moscow University, The Geological Institute (Moscow), The Industrial and Research Institute for Construction Engineering (Moscow), The Soil Institute (Pushino, Moscow Area) and others. ...
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