Y. geinitzi
View in CoL
with
Temnothorax gracilis ( Mayr, 1868)
,
Y. geinitzi
with
Temnothorax
sp. nov.,
Oecophylla brischkei Mayr, 1868
View in CoL
with
L. schiefferdeckeri
View in CoL
( Fig. 1
View Fig
), and
Bradoponera meyeri Mayr, 1868
with
F. flori
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. Two contain only Holarctic species: a
F. flori
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with
L. schiefferdeckeri
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and
Myrmica
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sp.n. with
Plagiolepis kuenowi Mayr, 1868
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. One contains only tropical ants:
Gnamptogenys europea ( Mayr, 1868)
with
N. pygmaea
View in CoL
.
As in the syninclusions with
Ctenobethylus
View in CoL
reported by Wheeler (1915) (see above), four pieces of amber contain
C. goepperti
View in CoL
with Holarctic species: three with
L. schiefferdeckeri
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and one with
F. flori
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, but three include
C. goepperti
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with the tropical species
N. pygmaea
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,
Tetraponera simplex ( Mayr, 1868)
View in CoL
and
Dolichoderus passaloma Wheeler, 1915
. Two syninclusions with
Monomorium pilipes Mayr, 1868
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contain tropical genera:
Tapinoma electrinum Dlussky, 2002
View in CoL
and
Y. geinitzi
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, and one of
M. mayrianum Wheeler, 1915
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is with
F. flori
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and another one is with
C. goepperti
View in CoL
.
Thus, regardless of whether
Ctenobethylus
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and
Monomorium
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are considered Holarctic or tropical, they occur in syninclusions with species of both groups, which not only emphasizes the mixed nature of the amber fauna, giving strong support for the cooccurrence of tropical and Holarctic species at the same time and place, and these syninclusions are no less important than those containing definitely Holarctic and tropical species.
The ratio of Holarctic and tropical taxa is important for a better understanding the climatic conditions in amber forests and the age of amber biota as a whole.
After adding
Aphaenogaster
,
Temnothorax
,
C. mengei
,
Eocenomyrma
and
N. pygmaea
the ratio of Holarctic to tropical s.l. elements for representative collections is 2.8–3.5 for Baltic amber, 1.6 for Danish amber and 2.1 for both Bitterfeld and Rovno ambers ( Table 2). It is interesting that at least for two amber faunas with the same (Danish, 1.6) or very similar (Rovno, 1.9) ratios were calculated for biting midges (
Ceratopogonidae
): predominance of Holarctic biting midges in the representative collections from other ambers is even higher than for Holarctic ants ( Perkovsky, 2017).
In the early Eocene at Messel and middle Eocene at Eckfeld, tropical taxa absolutely dominate, nearly as much as in tropical climate Oligocene Sicilian amber ( Emery, 1891; Wappler, 2003; Dlussky et al., 2008, 2009; Archibald et al., 2011; Dlussky, 2012; Dlussky, Wedmann, 2012). The prevalence of Holarctic species in all representative amber ant collections is associated with decreasing of temperature in the late Eocene (see below). The ratio of Holarctic and tropical specimens in the former private amber ant collection of M. Kutscher from Bitterfeld amber (now in the collection of Geowissenschaftlicher Zentrum der Georg-August-Universität Göttingen) is only 0.98, but it is highly biased, containing mainly rare and ‘exotic’ ants ( Dlussky, Rasnitsyn, 2009).