An Eocene bee in Rovno amber, Ukraine (Hymenoptera, Megachilidae)

2006·0.65 MB·English

by  Engel, Michael S

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JSIovitates
PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY
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Number 3506, 12 pp., 2 figures January 12, 2006
An Eocene Bee in Rovno Amber, Ukraine
(Hymenoptera: Megachilidae)
MICHAEL S. ENGEL1 AND EVGENY E. PERKOVSKY2
CONTENTS
Abstract . 2
Introduction . 2
Material and Methods . 3
Systematic Paleontology . 3
Ctenoplectrella zherikhini, new species. 3
Key to species of Ctenoplectrella. 4
Discussion . 4
Acknowledgments . 8
References . 8
Appendix 1: Fossil Megachilidae . 10
1 Division of Invertebrate Zoology, American Museum of Natural History; Division of Entomology (Paleoento-
mology). Natural History Museum, and Department of Ecology & Evolutionary Biology, Snow Hall, 1460 Jayhawk
Boulevard, University of Kansas, Lawrence, Kansas 66045-7523 ([email protected]).
2 Schmalhausen Institute of Zoology, National Academy of Sciences of Ukraine, 15 Bogdana Khmel’nitsky Str.,
Kiev 30, 01-601 Ukraine.
Copyright © American Museum of Natural History 2006 ISSN 0003-0082
2 AMERICAN MUSEUM NOVITATES NO. 3506
ABSTRACT
A new species of fossil bee is described and figured in Late Eocene amber from the Rovno
region of the Ukraine. Ctenoplectrella zherikhini n.sp. (Megachilinae: Ctenoplectrellini), is
similar to those species known from the slightly older Baltic amber (middle Eocene). This is
the first bee discovered in the Rovno amber and the fourth species for its genus. The presently
known megachilid bee fossils are summarized and the geological history of the family re¬
viewed.
INTRODUCTION ing deposits is referred to the northwestern
margin of the Ukrainian Crystalline Shield.
Bees are among the most recognizable in¬
Here terrestrial and shallow, marine, Paleo¬
sects and have endeared themselves to hu¬
gene sediments along with terrestrial (mainly
mans owing to their diligent pollination ser¬
bog-lacustrine) Neogene and predominantly
vices (agricultural and otherwise) and, in the
Anthropogene glacial formations overlie the
case of the honey bees, the honey and wax
Pre-Cambrian layers and their weathering
they produce. Those species of the family
surface. The Paleogene deposits in the most
Megachilidae are most familiar for the leaf-
complete sections contain the Buchak (Lu¬
cutter bees, whose semicircular swaths from
tetian), Kiev (Bartonian), Obukhov (Priabon-
petals and leaves are the frustration of gar¬
ian), Mezhigorje (Rupelian), and Berek
deners. Although our knowledge of the bees
(Chattian) Formations. Amber occurs in al¬
is advanced by comparison to most other lin¬
most all stratigraphic units of the sedimen¬
eages of Hymenoptera, understanding of
tary cover, although it has not been found in
their evolutionary history is compromised by
the Buchak deposits and is extremely rare in
the paucity of information from the fossil
those of the Kiev. However, even in the part
record. Whereas wasps and ants are frequent¬
of the section where amber occurrences are
ly plentiful in Cenozoic or Cretaceous de¬
abundant, its content in the rock is substan¬
posits, bees are scarcely recovered despite
tially different between layers. The richest
their almost assured presence and perhaps
placers are associated with the Obukhov
abundance in the same paleo-habitats. The
(Late Eocene) and Mezhigorje (Early Oli-
history of bees is partially compromised by
gocene) Formations. The Obukhov and Me¬
the poor preservation of organisms in arid
zhigorje Formations seem to have been
habitats, since it is in such environments that
bees, particularly basal bee lineages, flourish. formed in shallow water zones of marine ba¬
This rarity makes the discovery of any bee sins, with deep water areas situated in the
fossil of importance and excitement. Herein Pripyat and Dnieper-Donetsk depressions.
we provide the description of an Eocene bee The shorelines of these basins lay within the
preserved as an amber inclusion from the Ukrainian Shield. The zone of littoral shal¬
Rovno region of the Ukraine. This is the first low waters in this shield was apparently the
bee recovered from these deposits (Klesov area where the formation of amber placers in
and Dubrovitsa deposits of the Obukhov For¬ the sea was simultaneous with the accumu¬
mation) and was found in the collection of lation of primary sediment material of latter
Rovno amber inclusions acquired by the In¬ formations. The Klesov deposit is often mis¬
stitute of Zoology of the National Academy interpreted as of Oligocene age (e.g., Tutskij
of Sciences of Ukraine in Kiev. The fossil is and Stepanjuk, 1999; Weitschat and Wichard,
a representative of Ctenoplectrella, an extinct 2002). The source of this confusion stems
genus of megachiline bees previously known from Maidanovich and Makarenko (1988)
only from Baltic amber (Engel, 2001). To the wherein in one section the Klesov and Dub¬
description we have appended an account of rovitsa are referred to the Oligocene whereas
the geological history of the Megachilidae in all other sections of the same paper they
(vide etiam Engel, 1999) and an updated are correctly assigned to the Late Eocene.
checklist of the known fossils for the family. Most inclusions from the Institute of Zo¬
Structurally, the region of the amber-bear¬ ology are from the Klesov deposit. A list of
2006 ENGEL AND PERKOVSKY: UKRAINIAN FOSSIL BEE 3
Fig. 1. Photomicrograph of Ctenoplectrella zherikhini n.sp. (holotype female, UA-1748). Specimen
length 5.2 mm.
the 16 insect orders and 97 families reported for flagellomere, metasomal sternum, and
from Rovno amber was provided by Perkov- metasomal tergum, respectively. Measure¬
sky et al. (2003, to which can be added the ments were made with an ocular micrometer
Aleyrodidae and Megachilidae). Although on an Olympus SZX12 stereomicroscope.
roughly contemporaneous with the more his¬ All metrics should be considered approxi¬
torically studied Baltic amber, the fauna of mate since the optimal visual angle was not
Rovno amber is quite distinct. More than always achievable owing to the state of pres¬
26% of the ants represent genera and species ervation. Photomicrographs were prepared
not found in Baltic amber (Dlussky and Per- using a Microptics ML-1000 Digital Imaging
kovsky, 2002) and all of the more than 50 System. Comparative material of Baltic am¬
species of gall midges are unique to Rovno ber species of Ctenoplectrella was studied by
amber (Fedotova and Perkovsky, 2004, 2005; the senior author in the collections of the
Perkovsky and Fedotova, 2004). In particu¬ American Museum of Natural History (New
lar, the Rovno amber fauna differs from that York) and the University of Kansas Natural
of the Baltic by the dominance of Sciaridae History Museum (Lawrence).
and other forest litter Diptera, whereas Chi-
ronomidae and other aquatic insects are un¬ SYSTEMATIC PALEONTOLOGY
derrepresented (Perkovsky et al., in prep.).
Ctenoplectrella zherikhini, new species
MATERIAL AND METHODS Figure 1
Morphological terminology and format for Diagnosis: The new species is most sim¬
the description follow that of Engel (2001). ilar to C. grimaldii Engel in Baltic amber as
The abbreviations F, S, and T are employed both have the pleura impunctate and the im-
4 AMERICAN MUSEUM NOVITATES NO. 3506
bricate propodeum and metasoma. Cteno- with abundant, moderate length to long setae,
plectrella zherikhini, however, differs by the setae with short branches. Terga with sparse,
fuscous setae on the inner surfaces of the tar¬ short, simple, appressed or suberect setae, se¬
si; long, branched setae of the propodeum; tae longer and suberect to erect on lateral bor¬
and tapering gena. ders (some setae laterally with a few, minute
Description; Female. Total body length branches); sternal scopa composed of dense,
approximately 5.2 mm. Head apparently long, erect, simple setae.
slightly wider than long; face relatively flat; Holotype: Female, Rovno amber, UA-
gena much narrower than compound eye in 1748 (fig. 1), labeled “Holotype, Ctenoplec-
lateral view, gena tapering slightly in width trella zherikhini Engel & Perkovsky [red la¬
(widest dorsally, narrower ventrally). Basal bel]”. Deposited in the Institute of Zoology,
vein confluent with cu-a; second abscissa Rs National Academy of Sciences of Ukraine,
basad lm-cu by approximately six times vein Kiev.
width; other forewing venational details un¬ Etymology: The specific epithet is a pat¬
observable (forewing is obliquely torn); eight ronymic for the late Dr. Vladimir V. Zheri-
distal hamuli, arranged in a single, evenly khin (1945-2001). Dr. Zherikhin was a
spaced series. prominent coleopterist and paleoentomolo-
Outer surface of mandible impunctate and gist whose untimely death has deprived us of
smooth. Clypeus with small punctures sepa¬ his unrivaled scholarship and warm humor.
rated by a puncture width or less, integument
between smooth. Supraclypeal area appar¬ Key to Species of Ctenoplectrella
ently sculptured as on clypeus. Integument of (modified from Engel, 2001)
face mostly obscured by thin layer of Schim-
1. Crossvein 2rs-m strongly and doubly arcuate,
mel and debris toward vertex. Gena with mi¬
thus second submarginal cell more strongly
nute punctures separated by about a puncture produced toward wing apex along its posterior
width or less, integument between smooth. margin; medioapical margin of clypeus straight
Notal integument obscured by debris. Pleura
impunctate and smooth. Lateral surface of — Crossvein 2rs-m relatively straight and there¬
propodeum (basal and posterior surfaces ob¬ fore second submarginal cell not more strongly
scured) impunctate and faintly imbricate. produced toward wing apex along its posterior
margin; medioapical margin of clypeus gently
Metasomal terga and sterna faintly imbricate
convex . C. viridiceps Cockerell
and impunctate.
2. Pleura impunctate; metasomal terga faintly im¬
Integument dark brown, without macula-
bricate .3
tions. Wing membrane hyaline; veins strong
—Pleura with coarse, faint punctures; metasomal
and black. Pubescence generally white or terga with small, scattered punctures .
off-white, except setae on inner surfaces of . C. cockerelli Engel
tarsi fuscous (dark brown to black). 3. Propodeal setae long, erect, and branched; tar¬
Mandible with minute, simple, appressed sal setae fuscous; gena tapering .
setae. Clypeus with minute, appressed, simple . C. zherikhini n.sp.
setae not obscuring integument; similar setae —Propodeal setae scattered, short, and simple;
tarsal setae white or off-white; gena of rela¬
also on face below level of antennal sockets,
tively equal width along its length .
intermixed with scattered, longer, erect setae
. C. grimaldii Engel
on face, clypeus, supraclypeal area, gena, and
postgena. Pleura with scattered, short, simple
DISCUSSION
setae, setae becoming gradually longer ven¬
trally. Lateral surface of propodeum with The historical record of Megachilidae is
abundant moderate length to long, branched relatively good, with the amount of material
setae. Pubescence of legs generally simple being second only to that of the Apidae.
and of moderate length although becoming From more recent history are Bronze Age
long on outer surfaces of tibiae and tarsi; in¬ remains of burrows perhaps originating from
ner surface of metafemur without pubescence, the activity of osmiines in Israel (Buxton,
outer, anterior, and posterior surfaces with 1932). True fossil material of the family (i.e.,
moderate length, branched setae; metatibia Pre-Quaternary) is presently confined to the
2006 ENGEL AND PERKOVSKY: UKRAINIAN FOSSIL BEE 5
Cenozoic and to deposits of the Northern Pararhophitinae, consisting of a single genus
Hemisphere (appendix 1). However, the zoo¬ with three species, are restricted to the des¬
geography of the modern lineages coupled erts of Central Asia, Asia Minor, Arabia, and
with the temporal and phylogenetic distri¬ northeastern Africa. The species of Parar-
bution of known fossils (and trace fossils) is hophites are small, sparsely setose, mostly
quite revealing, suggesting at least a prelim¬ yellowish bees that are not commonly en¬
inary outline of megachilid evolutionary his¬ countered. The genus is clearly relic and
tory. united to the Fideliinae by a unique cocoon
The almost complete lack of fossils from construction by the mature larvae (in both
the Cretaceous limits any discussion of early subfamilies the larvae ingest sand and then
bee evolution. The oldest fossil of a bee is defecate in thin strips running from the poles
Cretotrigona prisca (Michener and Grimal¬ of the cocoon). The hdeliines are generally
di) from the Late Cretaceous of eastern North robust bees and atypical for the family in the
America (Michener and Grimaldi, 1988; En¬ densely setose legs, termination of the sub-
gel, 2000). Although Cretotrigona is, quite antennal sulci on the ventral margin of the
unfortunately, a singular fossil, it reveals antennal sockets, and three submarginal cells
much owing to its derived position in a de¬ (symplesiomorphies for the family), among
rived family (Apidae), implying that signifi¬ other traits. Species nest in the ground and
cant stages of bee evolution had already oc¬ are restricted to xeric regions mostly in
curred by the Late Cretaceous. Earlier evi¬ southern Africa but with isolated represen¬
dence of bee activity is suggested by pre¬ tatives in Chile (i.e., Neofidelia) and Moroc¬
sumed halictine nests of Cenomanian age co (i.e., Fidelia ulrikei Warncke). Modern Fi¬
from Arizona (Elliott and Nations, 1998), deliinae are likely relictual, and the subfam¬
again indicating that bees are older than this ily perhaps once had a much broader distri¬
epoch, perhaps as old as late in the Early bution with surviving remnants in the
Cretaceous to early in the middle Cretaceous regions where it is found today (Engel, 2002,
(Engel, 2001, 2004c). Even in the absence of 2004a). This would be in general accord with
any fossil megachilids, the simple presence patterns for other hymenopteran groups with
of an apid (the sister-group of Megachilidae) similar “Gondwanan” distributions but
in the Late Cretaceous indicates that the me¬ which were in fact distributed widely in the
gachilid lineage was also present during this Northern Hemisphere during the Cretaceous
period. Indeed, most clades today considered and until the early Neogene (e.g., Scoleby-
as families of the Apoidea were likely estab¬ thidae, Megalyridae). Unfortunately, para-
lished during the middle Cretaceous (Engel, rhophitines and hdeliines do not occur in
2001, 2004c; Grimaldi and Engel, 2005). By habitats that produce amber (i.e., subtropical
the middle of the Eocene epoch, the period to tropical environments) and compression
from which most bee lineages are first fossils would likely be almost impossible to
known, representatives of the principal fam¬ distinguish from other large bees (in the case
ilies are already present and are apparently of the hdeliines) or small, sparsely setose
not basal (Engel, 2001). apoids (in the case of the pararhophitines).
Certainly origins of the family Megachil¬ Although a fossil hdeliine or pararhophitine
idae are, like the beginnings of all bee fam¬ would indeed be signihcant, particularly if
ilies, obscure. The family is organized into from some northern deposit for the hdeliines,
four subfamilies (vide Engel, 2005); the Par- such material is unlikely to be recovered.
arhophitinae, Fideliinae, Lithurginae, and The Lithurginae have more the habitus of
Megachilinae. Among the numerous charac¬ typical megachilid bees and share with the
ters supporting the family, the most note¬ Megachilinae reduced metatibial scopae (ex¬
worthy is the development and use of a me- cept Protolithurgus), subantennal sulci meet¬
tasomal scopa for pollen transport. Phyloge¬ ing the antennal toruli along the outer mar¬
netic studies support the notion that para- gins, and the presence of only two submar¬
rhophitines and hdeliines are basal to a ginal cells (i.e., lrs-m is absent). Species nest
Lithurginae + Megachilinae clade (e.g., in rotten or dried wood, without brood cell
Roig-Alsina and Michener, 1993; fig. 2). The linings. The subfamily is noteworthy for the
6 AMERICAN MUSEUM NOVITATES NO. 3506
LITHURGINAE
F1DELIINAE / MEGACHI LINAE
Fig. 2. Phylogeny of Megachilidae, with significant fossils indicated; note that Osmiini is likely
paraphyletic and Anthidiini includes Dioxyini. Oe = Oeningen; Sh = Shandong; DA = Dominican
amber; Ro = Rott; FI = Florissant; BA = Baltic amber (including the Ukrainian amber fossil); refer
to appendix 1. Arrows indicate the position and age of particularly important fossil records for Antho-
phila. Cretotrigona is currently the oldest body fossil of the lineage sister to Megachilidae (i.e., Apidae)
(Michener and Grimaldi, 1988; Engel, 2000). The principal extant lineages are represented by Neofidelia
2006 ENGEL AND PERKOVSKY: UKRAINIAN FOSSIL BEE 7
presence of the metabasitibial plate (repre¬ gachilidae in Lewis, 1969) was tantalizing
sented only posteriorly in modern taxa), the not only as an early record of this genus but
pygidial plate, metatibial spicules (in modern as one of the only parasitic bee fossils. The
taxa), and the shortened first metasomal ter- specimen, however, is preserved solely as a
gum, which is flattened in profile. The ear¬ forewing and cannot be assigned to any par¬
liest record of lithurgines is Protolithurgus ticular megachiline tribe, let alone to the par¬
ditomeus Engel in Baltic amber, a species asitic genus Coelioxys (Engel, 2004b). Cock¬
that is also sister to all other members of the erell (1909a) referred to a fossil Osmia from
subfamily (Engel, 2001). Cladistic study has Germany originally identified by Oswald
also suggested that the most basal, living Heer under the manuscript name ” Osmia kir-
members of the subfamily are those species byancT, but subsequently recognized the
of the genus Trichothurgus from southern specimen as a polistine wasp (Cockerell,
South America (Engel, 2001). Trichothurgus 1914; Cockerell did not cite the manuscript
consists of robust, densely setose bees rem¬ name in 1909a but did so in 1914). Most of
iniscent of fideliines but otherwise having the fossil Megachilinae need to be critically
typical lithurgine-megachiline traits. The de¬ revised.
rived genus Lithurgus consists of two prin¬ The famous leaf-cutter bees (Megachile
cipal lineages: Lithurgopsis in the New group I sensu Michener, 2000, or Megachile
World (Argentina to the northern United in the traditional sense, e.g., Michener, 1962;
States) and Lithurgus proper (native to Eur¬ Mitchell, 1980) are a derived group of the
asia, Africa, and Australia), which should Megachilini. This group is certainly older
perhaps be distinguished as separate genera than the middle Eocene. Although the earli¬
(vide Engel, 2001). The enigmatic Lithurgus est body fossils of this lineage are those from
rubricatus Smith from Australia is notable the Eocene-Oligocene boundary of Floris¬
for the occurrence of elongate arolia and sant, Colorado (appendix 1), the distinctive
should perhaps be transferred to its own sub¬ semicircular traces of their activity in fossil¬
genus. A putative Lithurgus was described ized leaves are known from the Early to mid¬
from the Miocene of Germany (Cockerell, dle Eocene of various deposits in North
1909a: appendix 1) but although certainly a American and Europe (e.g., Berry, 1916,
megachilid, the generic assignment is dubi¬ 1931; Brooks, 1955; Lewis, 1994; Wappler
ous. Thus, Protolithurgus is the only defini¬ and Engel, 2002, 2003; Wappler, 2003a,
tive fossil of the subfamily. It is likely that 2003b). Leaves with such cuts are also
the lithurgines originated in the Southern known from other Tertiary deposits (Cock¬
Hemisphere (i.e., of Gondwanan origin), dif¬ erell, 1908, 1910; Meyer, 2002), as well as
ferentiating from Megachilinae sometime in an inclusion in Dominican amber with a
the middle to Late Cretaceous. specimen of Sphaerodactylus (Grimaldi,
Of the megachilids, Megachilinae are the 1996). Interestingly, the leaf cuts from the
most numerous in the fossil record (appendix Eocene are presently the earliest evidence of
1), with each tribe represented to some extent the Megachilidae as a whole (fig. 2), being
(fig. 2). Early accounts of “Osmia”, “Chal- slightly older than the Baltic amber fossils.
icodoma”, and “Megachile” in Baltic amber The presence of derived Megachile in the
(e.g., Menge, 1856; Brischke, 1886; Bacho- Early Eocene, however, indicates that more
fen-Echt, 1949; Scudder, 1885, 1891) were basal divergences in the Megachilini must
likely specimens of Ctenoplectrella, which have taken place earlier in the Paleocene or
has the general habitus of an osmiine. Ref¬ Latest Cretaceous. Similarly, divergences be¬
erence to a fossil ” Coelioxys” sp. (Lewis et tween the tribes of Megachilinae and con¬
al., 1990; earlier referred to as simply Me- comitantly between the subfamilies must
<—
profuga Moure and Michener (Fideliinae + Pararhophitinae), Lithurgus chrysurus (Fonscolombe) (Li-
thurginae), Anthidium florentinum Latreille (Anthidiini), Hoplitis fulgida (Cresson) (Osmiini), and Me¬
gachile Candida Smith (Megachilini).
AMERICAN MUSEUM NOVITATES NO. 3506
have occurred even earlier (fig. 2). The ex¬ of bees, XXXIV. Annals and Magazine of Nat¬
ploration of Cretaceous and Paleocene fossil ural History, series 8 7: 225-237.
Cockerell, T.D.A. 1913. Some fossil insects from
locales will be of considerable importance
Florissant. Canadian Entomologist 45: 229-
for unraveling the earliest evolution of the
233.
Megachilidae, particularly amber sites. Al¬
Cockerell, T.D.A. 1914. Miocene fossil insects.
though fossils of bees are admittedly rather
Proceedings of the Academy of Natural Sci¬
uncommon, if not rare, in most places, the
ence, Philadelphia 66: 634-648.
remains of leaves are abundant and may con¬ Cockerell, T.D.A. 1917. New Tertiary insects.
tinue to provide data on the Cenozoic paleo- Proceedings of the United States National Mu¬
geographical and temporal distribution of the seum 52: 373-384.
leaf-cutting clade. Cockerell, T.D.A. 1923. Two fossil Hymenoptera
from Florissant (Vespidae, Megachilidae). En¬
tomological News 34: 270-271.
ACKNOWLEDGMENTS
Cockerell, T.D.A. 1925a. Descriptions and records
We are grateful to Dr. Vladimir Blagode- of bees, CVI. Annals and Magazine of Natural
rov and Dr. Torsten Wappler for their insight¬ History, series 9 16: 416-423.
ful comments on the manuscript. Partial sup¬ Cockerell, T.D.A. 1925b. Tertiary insects from
port for this study was provided by NSF EF- Kudia River, Maritime Province, Siberia. Pro¬
ceedings of the United States National Museum
0341724 (to MSE). This is contribution No.
68: 1-16.
3430 of the Division of Entomology, Natural
Dlussky, G.M., and E.E. Perkovsky. 2002. Ants
History Museum, University of Kansas.
(Hymenoptera, Formicidae) from the Rovno
amber. Vestnik Zoologii 36: 3-20. [In Russian]
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APPENDIX 1
Fossil Megachilidae poorly understood (e.g., Pictet, 1854, considered
it similar to Bombus\). A specimen of Heriades
The following is a checklist of described fos¬
sp. in African copal is also excluded since it is
sil bees of the family Megachilidae (updated and
likely of an extant species (Engel, 2001). The rec¬
modified from Engel, 1999). Note that although
ord of Megachile sp. in Baltic amber by Gerlach
in some instances assignment to family, subfam¬
(1989) was a misidentification with Ctenoplec-
ily, and tribe may be reasonable, many generic trella (Engel, 2001). The listing of Stelis senecio-
assignments for compression fossils must be taken phila Cockerell as a fossil from the Florissant
with caution. Apiaria dubia Germar (1849) of the Fossil Beds (Meyer, 2002) is erroneous as this is,
Oligocene of Germany is excluded despite its in¬ in fact, a living species that was simply described
clusion in Osmia by Giebel (1856; vide etiam from near modem Florissant, Colorado, and in the
Scudder, 1891; Handlirsch, 1907) since it is so same paper with some actual fossil taxa.
Taxa References
EOCENE
Baltic Amber (Lutetian)
Protolithurgus ditomeus Engel Engel, 2001
Ctenoplectrella cockerelli Engel Engel, 2001
Ctenoplectrella grimaldii Engel Engel, 2001
Ctenoplectrella viridiceps Cockerell Cockerell, 1909a
= Ctenoplectrella dentata Salt, 1931
= Ctenoplectrella splendens Kelner-Pillault, 1970