Turkish Journal of Zoology
http://journals.tubitak.gov.tr/zoology/
Research Article
Turk J Zool
(2014) 38: 444-459
© TÜBİTAK
doi:10.3906/zoo-1309-5
Distribution data on the family Melittidae (Hymenoptera) of Turkey
with considerations about their importance as pollinators
Hikmet ÖZBEK*
Department of Plant Protection, Faculty of Agriculture, Atatürk University, Erzurum, Turkey
Received: 13.10.2013
Accepted: 10.01.2014
Published Online: 20.05.2014
Printed: 19.06.2014
Abstract: The examination of specimens collected from various parts of Turkey, mainly from East Anatolia, and an overview of the
literature allows us to reach the conclusion that the Melittidae of Turkey are represented by 3 genera, Dasypoda (14 species), Macropis (2
species), and Melitta (9 species), with a total of 25 species including Dasypoda toroki Michez, 2004, a new record for the Turkish fauna.
Dasypoda hirtipes (Fabricius, 1793), Melitta (Melitta) leporina (Panzer, 1799), and M. dimidiata Morawitz, 1876 are the most widespread
and abundant species. Dasypoda friesana Schletterer, 1890; D. longigena Schletterer, 1890; D. patinyi Michez, 2002; D. visnaga (Rossi,
1790); Melitta rasmonti Michez, 2007; M. nigricans Alfken, 1905; M. singular Michez, 2012; M. haemorrhoidalis; and M. melanura
(Nylander, 1852) are very rare species and most are known from one locality only. The rest of the species are moderately distributed
in Turkey. Five species of Dasypoda and 3 species of Melitta have type localities from Anatolia; of these, D. longigena, M. bicollaris,
M. singular, M. rasmonti, and D. warnckei are endemic to Anatolia. New distribution areas were added for each species and the plant
species visited were included. Although the genus Dasypoda visits mostly wild flowers, D. hirtipes also frequently visits sunflowers and it
is an important pollinator of this plant. In the case of the genus Macropis, females collect oil and pollen from the flowers of Lysimachia
(Primulaceae) as larval food. Melitta leporina, M. dimidiata, and M. bicollaris are important pollinators of fodder crops, such as alfalfa,
sainfoin, and various clover species and some other legumes.
Key words: Hymenoptera, Melittidae, Dasypoda, Macropis, Melitta, fauna, new record, pollinator, Turkey
1. Introduction
Bees are in the Apiformes group of Apoidea (Hymenoptera),
which is a diverse group of pollen-eaters comprising
approximately 18,000 described species (Michener, 2007;
Michez et al., 2009). They occur in most ecosystems
where they are usually the most important pollinators,
and therefore they play a key role in ecosystem function
(O’Toole, 1993; Waser and Ollerton, 2006). Currently, 9
bee families are recognized, and Melittidae is one of the
smallest bee families with more than 200 species in 14
genera (Engel, 2005; Michener, 2007; Michez et al., 2009,
2012).
The family Melittidae includes short-tongued bees.
Michener (2007) emphasized that no single character is
unique to Melittidae, although the combination of a short,
pointed glossa and unspecialized first 2 segments of the
labial palpus with a slender, V-shaped lorum; tapering
mentum; and elongate, fully exposed middle coxa as in
long-tongued bees separates the Melittidae from other
short-tongued bee families.
*Correspondence: hozbek@atauni.edu.tr
444
Members of this rather small family are found
primarily in temperate regions of the northern hemisphere
and Africa, the greatest number of genera and species
being found in warm xeric areas. Maximal diversity
occurs in southern Africa, where all 3 subfamilies occur.
The Palearctic region also supports a moderate number
of species and genera. In the Nearctic region the family is
rare, unknown in tropical America and Australia, as well
as in the Indo-Malayan area (Michener, 2007; Michez et
al., 2009).
Michener (1981) divided the Melittidae into 3
subfamilies:
Meganomiinae,
Dasypodainae,
and
Melittinae. Dasypodainae and Melittinae occur in Turkey
and Meganomiinae is known from Africa and Arabia only
(Michener, 1981), whereas, according to Engel (2001), the
Melittidae are subdivided into 4 subfamilies: Dasypodainae,
Macropidinae, Meganomiinae, and Melittinae.
The Dasypodainae are relatively species-rich,
comprising more than 100 species in xeric areas of both
the Old World and the Nearctic region (Michener, 2007;
Michez et al., 2009, 2010). Dasypoda is the only widespread
ÖZBEK / Turk J Zool
genus, which occurs from the temperate to the xeric areas
of the Palearctic, including Turkey. Dasypoda determines
the northern limit of the Dasypodainae to 62°N (Michez
et al., 2004a). The other Dasypodainae genera, Capicola
Friese, 1911; Eremaphanta Popov, 1940; Hesperapis
Cockerell, 1898; and Samba Friese, 1908, are each endemic
in different Old World and Nearctic semideserts (Michez
and Pauly, 2012). The Melittinae comprise 4 genera: Melitta
Kirby, 1802; Macropis Panzer, 1809; Rediviva Friese, 1911;
and Redivivoides Michener, 1981 (Michener, 1981). Melitta
and Macropis have Holarctic distributions; Melitta is the
most widespread and probably one of the most diverse,
while Rediviva and Redivivoides are endemic in Africa
(Michener, 2007; Michez and Eardley, 2007). Two genera,
Macropis and Melitta, occur in Turkey.
Some important studies have been conducted on
Melittidae in the Palearctic: Quilis (1928) reviewed the
Spanish species of Dasypoda. The West Palearctic region
of Melittidae was studied by Warncke (1973), who listed
17 Melitta species including 4 new species and several
subspecies, 3 Macropis species, and 18 Dasypoda species
including 1 new species and several subspecies. The
Melittidae species of China and Dasypoda species of
European Russia were studied by Wu (1978) and Radchenko
and Pesenko (1989), respectively. An annotated account of
the species of Dasypoda was provided by Baker (2002), who
described D. litigator from Iran. Michez (2002) described
D. patinyi from Syria. In a phylogenetic study, Michez et al.
(2004b) recognized 4 subgenera. Michez and Pauly (2012)
described D. riftensis from Ethiopia. This species was the
first record of the genus Dasypoda in sub-Saharan Africa.
Most of the genera in Melittidae were revised: Michez
et al. (2004a) revised the West Palearctic Dasypoda and
recognized 28 species, including 4 new species. Michez
and Eardley (2007) provided an overview of the literature
and their examination of many specimens from various
collections, including type materials, enabled the first
monographic revision of the genus Melitta. They listed 43
species, including 2 new ones in the 2 subgenera Melitta s.
str. and Cilissa. More recently, Michez et al. (2012) described
4 Melitta species. Michez and Patiny (2005) revised the oilcollecting bee genus Macropis Panzer and described a new
species from Laos. Other revisional studies included those
on Capicola Friese (Michez et al., 2007a), Eremaphanta
Popov (Michez and Patiny, 2006), Hesperapis Cockerell
(Stage, 1966; Michener, 1981), Promelitta Warncke
(Michez et al., 2007b), Meganomia Cockerell (Michener,
1981), and Rediviva Friese (Whitehead et al., 2008).
Studies on the family Melittidae in Turkey are very
restricted. Warncke (1973) presented a catalog and key
for 38 West Palearctic species. Among them, 15 species (1
Macropis, 8 Dasypoda, and 6 Melitta species) were indicated
to occur in Turkey. He described Melitta bicollaris from
Erzurum, Dasypoda pyriformis tubera from İçel (Mersin),
and D. braccata anatolica from Konya. Recently, additional
species were described from Turkey: Dasypoda warnckei
Michez, 2004 from Kayseri; Melitta rasmonti Michez,
2007 from Hakkari; and M. singular Michez, 2012 from
Nevşehir (Michez et al., 2004a, 2012; Michez and Eardley,
2007). Local researcher Özbek (1976, 1979, 1980, 2008,
2011) highlighted the importance of various Melitta
species on the pollination of different fodder crops, such
as alfalfa, sainfoin, and red clover.
The aim of this paper is to present the latest knowledge of
the Turkish Melittidae bees, their geographic distribution,
plant species visited, and biogeographical affinities. The
present contribution is based on material collected from
various parts of Anatolia by the author and his colleagues
since the 1960s and literature data.
2. Materials and methods
Most of the bee specimens were collected by sweeping
with an insect net from various habitats in different
localities, mainly in eastern Anatolia, since the 1960s. The
catches of bees were made during the flowering periods of
plants, between April and September. In addition to bees,
the flowering plants that particular bee species visited
were noted or the samples were picked. Identification of
the bees was verified by comparison with the preserved
specimens, which were determined by the late K Warncke,
late YA Pesenko, and B Tkalcu at the Entomology Museum
of Erzurum, Turkey (EMET). Additionally, the keys
provided by Warncke (1973) and Michez et al. (2004a)
were followed. Certain undetermined specimens were
determined by D Michez. Host-plant data were based on
the field records and labels of pinned specimens of EMET.
The material mentioned in this study was deposited in the
collection of EMET, along with a few specimens deposited
in the collection of the American Museum of Natural
History (AMNH).
3. Results
The examination of the specimens collected from various
parts of the country and an overview of the literature allow
us to reach the conclusion that the Melittidae of Turkey are
represented by 3 genera, Dasypoda (14 species), Macropis
(2 species), and Melitta (9 species), with a total of 25
species.
3.1. Genus Dasypoda Latreille, 1802
The genus Dasypoda is characterized by the presence
of only 2 submarginal cells on the fore wing and a welldeveloped scopa of the female hind leg, black body, vertex
elevated, no basitibial plate (Michener 1981; Michez et al.,
2004a, 2004b). Michez et al. (2004b) described 4 subgenera
based on morphological cladistic analysis: Dasypoda s. st.;
Heterodasypoda Michez, 2004; Microdasypoda Michez,
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ÖZBEK / Turk J Zool
2004; and Megadasypoda Michez, 2004. The diversity
centers of each of the 4 subgenera are restricted to 1 of the
following parts of the Mediterranean region: the Balkans,
Morocco, and Spain (Michez and Pauly, 2012). Michez
(2005) listed 33 Dasypoda species that are common in the
Palearctic region from Portugal and the Canary Islands
to Japan, although most species are West-Palearctic and
especially abundant in the Mediterranean basin.
3.1.1. Dasypoda (Dasypoda) hirtipes (Fabricius, 1793)
Distribution: Whole of Europe; Asia from Turkey to
Mongolia and China; North Africa (Warncke, 1973; Wu,
1978; Michez et al., 2004a, 2004b). In Turkey: Amasya,
Balıkesir, Bursa, İstanbul, Kastamonu, Konya, Kütahya,
Nevşehir, Samsun, and Sinop (Warncke, 1973).
Material: Ağrı: Patnos, 28.07.1978, ♀, 15 ♂♂,
leg. H. Özbek, det. B. Tkalcu and H. Özbek. Ankara:
Şereflikoçhisar, 08.07.1998, 2 ♂♂, leg. C. Güçlü, det. D.
Michez and W. Thayse. Ardahan: Center, 11.08.1976, ♂,
leg. H. Özbek, det. B. Tkalcu; Posof, center, 28.08.1991, ♂,
leg. H. Özbek, det. H. Özbek; Sarıdarı, 28.08.1991, ♀, leg. E.
Yıldırım, det. H. Özbek; Türközü, 28.08.1991, 5 ♀♀, 3 ♂♂,
leg. H. Özbek, det. H. Özbek, D. Michez and W. Thayse.
Artvin: Yusufeli, İşhan, 17.09.1978, ♀, leg. H. Özbek, det.
B. Tkalcu; İşhan, 15.08.1991, 2 ♀♀, leg. E. Yıldırım, det.
H. Özbek; Sarıgöl, 700 m, 05.07.1994, ♂, leg. H. Özbek,
det. as D. altercator (Harris). Bilecik: Center, 600 m, ♂, leg.
E. Yıldırım, det. D. Michez and W. Thayse. Bitlis: Tatvan,
Nemrut Dağı, 23.07.2003, ♀, leg. H. Özbek, det. H. Özbek.
Erzincan: Çatalarmut, 09.08.1990, 3 ♂♂, leg. E. Yıldırım,
det. H. Özbek; Kemah, Alp, 19.07.1997, ♀, leg. H. Özbek,
det. H. Özbek; Üzümlü, Karakaya, 31.07.1991, 3 ♂♂, leg.
E. Yıldırım, det. Y. A. Pesenko as D. altercator; Tercan,
Yazıören, 29.06.1980, ♂, leg. H. Özbek, det. Y. A. Pesenko.
Erzurum: Atatürk University Campus, 12.07.1970, ♂, leg.
H. Özbek, det. K. Warncke; 24.07.1978, ♂, leg. H. Özbek,
det. B. Tkalcu; 18.08.1967, ♂ leg. O. Ecevit, det. H. Özbek;
20.08.1966, ♂, leg. H. Özbek, det. K. Warncke; 22.08.1968,
♂, leg. H. Özbek, det. H. Özbek; 01.08.1990, ♀, leg. K.
Gümüş, det. H. Özbek; Eşkinkaya, 20.07.2003, ♀, ♂, leg.
J. G. Rozen, H. Özbek, det. J. G. Rozen (American Natural
History Museum, AMNH); Palandöken, 2100 m, ♂, leg. E.
Yıldırım, det. H. Özbek; Horasan, Karakoçan, 19.07.1990, 3
♂♂, leg. E. Yıldırım, det. H. Özbek; Köprüköy, 26.08.1991,
♀, ♂ leg. H. Özbek, det. H. Özbek, D. Michez and W.
Thayse; Köprüköy, 19.07.1990, 2 ♂♂, leg. E. Yıldırım, det.
Y. A. Pesenko as D. altercator; Oltu, center, 20.07.1976, ♂,
leg. H. Özbek, det. B. Tkalcu; 14–16.09.1978, 2 ♂♂, leg.
H. Özbek, det. B. Tkalcu; Anzav deresi, 31.08.1996, ♀, ♂,
leg. G. Tozlu, det. H. Özbek; Ayvalı, 27.07.1990, ♂, leg. E.
Yıldırım, det. Y. A. Pesenko as D. altercator; 26.08.1993,
♀, leg. G. Tozlu, det. H. Özbek; Başaklı, 01.09.1978, 9
♂♂, leg. H. Özbek, det. Y. A. Pesenko as D. altercator;
07.09.1978, 4 ♂♂, leg. H. Özbek, det. H. Özbek; Başaklı,
446
14.09.1978, ♀, leg. H. Özbek, det. H. Özbek; Başaklı,
01.09.1988, 4 ♀♀, leg. H. Özbek, det. H. Özbek; Kayaaltı,
800 m, ♂, leg. G. Tozlu, det. D. Michez and W. Thayse;
Olur, Coşkunlar, 12.07.1991, ♂, leg. H. Özbek, det. H.
Özbek; Pasinler, center, 12.07.1980, ♂, leg. H. Özbek, det.
B. Tkalcu; 01.08.1983, ♂, leg. H. Özışık, det. D. Michez
and W. Thayse; Rabat, 2200 m, 13.06.1996, ♀, leg. H.
Özbek, det. H. Özbek; Şenkaya, Gaziler, 1650 m, 2 ♀♀,
leg. R. Kotan, det. H. Özbek; Tortum, Aksu, 16.09.1977, ♀,
leg. H. Özbek, det. H. Özbek, Aksu, 1450 m, 19.07.2003,
♂, ♀, leg. S. Çoruh, det. H. Özbek, det. D. Michez and W.
Thayse; Bağbaşı, 20.09.1978, 41 ♂♂, 4 ♀♀, leg. H. Özbek,
det. B. Tkalcu and H. Özbek; Şenkaya, Turnalı, 03.09.1990,
♀, ♂, leg. E. Yıldırım, det. H. Özbek. Eskişehir: Sivrihisar,
13.08.1990, ♀, leg. H. Özbek, det. H. Özbek. Iğdır: Aralık,
02.08.1997, ♂, leg. B. Gül, det. D. Michez and W. Thayse.
İzmir: Menemen, 900 m, 17.06.1994, ♂, leg. H. Özbek, det.
Michez and W. Thayse. Kars: Digor, 1770 m, 17.07.2005,
leg. H. Özbek, det. D. Michez and W. Thayse; Sarıkamış,
11.08.1983, 2 ♂♂, leg. H. Özbek, det. Y. A. Pesenko and
H. Özbek. Kayseri: Develi, 25.06.1980, 2 ♀♀, 3 ♂♂, leg.
H. Özbek, det. H. Özbek. Manisa: Alaşehir, 23.07.1990, 2
♂♂, ♀, leg. H. Özbek, det. H. Özbek. Samsun: 27.08.1981,
2 ♂♂, leg. O. Ecevit, det. H. Özbek. Sivas: Suşehri,
07.07.1999, 2 ♂♂, leg. H. Özbek, det. H. Özbek. Tokat:
Center, 19.08.1992, 3 ♀♀, leg. H. Özbek, det. H. Özbek;
900 m, 19.08.1992, ♂, ♀, leg. H. Özbek, det. D. Michez and
W. Thayse. Van: Edremit, 29.07.1978, 15 ♀♀, 7 ♂♂, leg. H.
Özbek, det. B. Tkalcu and H. Özbek; Gevaş, 29.07.1978, ♀,
leg. H. Özbek, det. H. Özbek; Kuzgunkıran, 29.07.1978, ♀,
leg. H. Özbek, det. H. Özbek. Yozgat: Yerköy, 09.08.1989,
3 ♂♂, leg. H. Özbek, det. H. Özbek.
Plants associated: Sunflower (Helianthus annuus L.)
and safflower (Carthamus tinctorius L.) are important
cultivated plant species that are frequently visited by D.
hirtipes. Wild plants: Apiaceae: Ammi visnaga (L.) Lam.,
A. majus L., Daucus carota L., Eryngium billardierei
F.Delaroche, E. bithynicum Boiss., E. creticum Lam.,
Ferula orientalis L., Heracleum spp., Pimpinella spp.;
Asteraceae: Achillea millefolium L., Arctium lappa L.,
A. minus (Hill), Carduus acanthoides L., C. nutans L.,
Centaurea drabifolia Sm., C. glastifolia L., C. iberica Trevir.
ex Spreng., C. hyalolepis Boiss., C. macrocephala Muss.
Puschk., C. solstitialis L., Cichorium intybus L. (the most
favorable plant), Cirsium arvense (L.), C. lappaceum
(M.Bieb), Echinops galaticus Freyn, E. orientalis Trautv.,
Onopordum acanthium (L.), O. illyricum L., O. tauricum
Willd., Pallenis spinosa (L.), Senecio spp., Taraxacum
aleppicum Dahlst., T. phaleratum G.E.Haglund ex Rech., T.
scaturiginosum G.E.Haglund, Tragopogon aureus M.Bieb.;
Convolvulaceae: Convolvulus arvensis L.; Dipsacaceae:
Cephalaria gigantea (Ledeb.), C. lycica Matthew, C. procera
Fisch. & Avé-Lall.; Euphorbiaceae: Euphorbia orientalis
ÖZBEK / Turk J Zool
L., Dipsacus laciniatus L., Scabiosa caucasica M.Bieb.;
Fabaceae: Melilotus officinalis L.; Lamiaceae: Salvia
aethiopis L., S. candidissima Vahl, S. multicaulis Vahl, S.
sclarea L., S. verticillata L., S. viridis L., Satureja sp., Stachys
annua L., S. cretica L., Thymus comptus Friv., T. leucostomus
Hausskn. & Velen., T. sipyleus Bois.; Scrophulariaceae:
Verbascum spp.
Remark: Dasypoda hirtipes is the most widespread and
abundant species of the genus in Turkey (Figure 1), as well
as outside of Turkey.
3.1.2. Dasypoda (Dasypoda) pyriformis Radoszkowski,
1887
Distribution: Italy, Balkan Peninsula, Croatia, Macedonia,
Albania, Bulgaria, Greece, Turkey (Michez et al., 2004a).
Michez (2012a) noted that D. pyriformis is present in the
northeastern part of the Mediterranean basin. Most of the
records are found in Greece. In Turkey: Denizli (Warncke,
1973); Amasya, as D. schlettereri Friese, 1900, which is a
synonym of D. pyriformis (Baker, 2002).
Material: Erzurum: Oltu, Başaklı, 01.08.1977, 2
♀♀ leg. H. Özbek, det. Y. A. Pesenko. Kars: Kağızman,
Kuloğlu, 1600 m, 01.08.2002, ♂, leg. M. Kesdek, det. D.
Michez and W. Thayse.
New localities: Erzurum and Kars.
Remark: Dasypoda pyriformis is a rare species
occurring in Turkey. It is a Mediterranean species; among
Asian countries, it occurs only in Turkey (Amasya). In the
present study it has been recorded from Erzurum and Kars
provinces, and Kars is its easternmost distribution record,
both in Turkey and the world.
Plants associated: Carduus sp., Centaurea glastifolia L.,
Cichorium intybus.
D. pyriformis
D. tubera
D. warnckei
D. pyrotrichia
D. argentata
D. braccata
D. friesana
D. longigena
D. patinyi
D. spinigera
D. aurata
D. toroki
D. visnaga
D. hirtipes
3.1.3. Dasypoda (Dasypoda) tubera Warncke, 1973
Distribution: Syria and Turkey. In Turkey: İçel and Hatay
(Warncke, 1973); Çukurova (rather abundant), Amasya,
Bilecik, Çorum, Eskişehir, İçel, İzmir, Hakkari, Hatay,
Taurus Mts. (Michez et al., 2004a).
Material: Adana: Akvatan, 22.05.1995, 2 ♂♂, leg. G.
Çankaya, det. D. Michez, W. Thayse and H. Özbek. Antalya:
Beydağları, Saklıkent yolu, Alimin Pınarı, 09.07.1991,
2 ♀♀, leg. H. Özbek, det. D. Michez, W. Thayse and H.
Özbek. İzmir: Menemen, 900 m, 17.06.1994, 2 ♀♀, ♂, leg.
H. Özbek, det. D. Michez, W. Thayse and H. Özbek.
Remark: Warncke (1973) described D. tubera from İçel
(Mersin) as the subspecies of D. pyriformis on the basis of
the male. Later, Michez et al. (2004a) found females and
additional males and described the male, and the status of
D. tubera was revised at the species level.
Plants associated: Carduus sp., Centaurea solstitialis,
and Cichorium intybus.
3.1.4. Dasypoda (Dasypoda) warnckei Michez, 2004
Distribution: Turkey. Kayseri (type locality), Niğde,
Nevşehir, Çorum, Denizli, Hakkari, Kars (Michez at al.,
2004a).
Material: Iğdır: Bayraktutan, 40°00′949N, 43°55′105E,
17.08.2005, ♀, ♂, leg. H. Özbek, det. D. Michez and W.
Thayse. Kars: Digor, 1770 m, 17.07.2005, ♂, leg. H. Özbek,
det. H. Özbek.
Plants associated: Cichorium intybus, Onopordum sp.
Remark: As far as is known, D. warnckei is endemic
to Turkey. It is not abundant, although it is a rather
widespread species from east to west (Figure 1). However,
it is remarkable to note that although I have made
intensive collections within Erzurum Province, I have not
encountered this species so far in Erzurum.
Mersin
Figure 1. Distribution of the genus Dasypoda in Turkey.
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ÖZBEK / Turk J Zool
3.1.5. Dasypoda (Heterodasypoda) pyrotrichia Förster,
1855
Distribution: Portugal, Spain, France, Greece, Bulgaria,
Macedonia, Israel, Syria, and Turkey (Michez et al., 2004a).
In Turkey: D. pyrotrichia was described from Anatolia
(Kleinasien) (Baker, 2002). It has already been recorded
in Erzurum, Gümüşhane, Kars, Kayseri, and Trabzon
(Michez et al., 2004a).
Material: Bayburt: Yaylapınar, 26.07.1992, ♀, leg. H.
Bostan, det. H. Özbek. Erzincan: Cevizli, 26.07.1993, 2
♀♀, leg. H. Özbek, det. K. Warncke and Y. A. Pesenko.
Erzurum: Atatürk University Campus, 10.06.2000, ♀,
leg. M. Kestek, det. H. Özbek; Ilıca, Ovacık, İspir road,
08.08.1990, ♂, leg. Ü. Avcı, det. D. Michez and W. Thayse;
Köşkköyü, 14.07.1992, ♂, leg. H. Özbek, det. D. Michez
and W. Thayse; Köprüköy, Örentaş, 04.07.2004, ♀, leg.
H. Özbek, det. H. Özbek; Oltu, Başaklı-Tutmaç border,
1700–2000 m, 02.07.2000, 15 ♀♀, 3 ♂ ♀, leg. H. Özbek
and Ö. Çalmaşur, det. H. Özbek; Kaleboğazı, 28.07.1990,
♂, leg. E. Yıldırım, det. H. Özbek; Pasinler, Çalıyazı,
13.07.1997, 9 ♂♂, leg. E. Yıldırım, det. H. Özbek; Rabat,
2400 m, 11.07.1996, 46 ♂♂, leg. E. Yıldırım, det. H. Özbek;
Pelitli, 14.07.1996, 2200 m, 14.07.1996, 2 ♀♀, 3 ♂♂, leg.
E. Yıldırım, det. H. Özbek; Şenkaya, Turnalı, 1800 m,
19.06.1991, ♀, leg. E. Yıldırım, det. D. Michez and W.
Thayse; 06.07.1990, ♀, leg. E. Yıldırım, det. H. Özbek;
Tortum, Aşağı Meydanlar, 04.07.1991, ♀, leg. H. Özbek,
det. D. Michez and W. Thayse; Doruklu Yaylası, 2000 m,
03.07.1991, ♀, 2 ♂♂, leg. H. Özbek, det. H. Özbek. Kars:
Sarıkamış, İssisu, 07.07.1991, 2 ♂♂, leg. E. Yıldırım, det.
H. Özbek, det. D. Michez and W. Thayse.
Plants associated: The samples were collected mainly
from the edges of wooded areas on Helianthemum
nummularium (L.) Mill. (Cistaceae). A few samples were
collected from Onopordum sp., Cirsium sp., Cephalaria
procera, Melilotus alba, and Salvia spp. Michez et al. (2008)
mentioned that D. pyrotrichia appears strictly oligolectic
on Cistaceae, although the sample was too small to reach a
definite conclusion.
3.1.6. Dasypoda (Megadasypoda) argentata Panzer, 1809
Distribution: It is widespread, particularly in western
Europe, Spain, France, Austria, Germany, Switzerland,
Italy, Poland, Bohemia, Slovakia, Croatia, Hungary,
Romania, Lithuania, Armenia, Azerbaijan, European
Russia, Macedonia, Sweden, Bulgaria, Greece, Ukraine,
Caucasus, Georgia. In Asia: Turkey, Kazakhstan, India,
Iran, Iraq, and Syria. In Africa: Algeria, Egypt, Tunisia
(Warncke, 1973; Michez et al., 2004a, 2004b). In Turkey:
Bayburt, Erzurum, Gümüşhane, İçel (Warncke, 1973);
Hakkari, Erzincan, Sivas (Michez et al., 2004a).
Material: Ağrı: Tahir Mt., 25.07.1978, ♂, leg. H.
Özbek, det. Y. A. Pesenko. Ardahan: Göle, 10.08.1977, 5
♂♂, leg. Özbek, det. Y. A. Pesenko and H. Özbek; 3 ♂♂,
448
leg. H. Özbek, det. Y. A. Pesenko. Bayburt: Demirdöven,
30.07.1992, ♀, leg. H. Özbek, det. H. Özbek; Kopdağı,
10.08.1991, 2 ♀♀, 2 ♂♂, leg. H. Bostan, det. D. Michez
and W. Thayse; Kopdağı, 2400 m, 13.08.1977, 4 ♀♀, leg.
Ö. Çalmaşur, det. H. Özbek. Bingöl: Solhan, Bağlan, 2122
m, ♂, leg. M. Kesdek, det. H. Özbek. Erzincan: Kızıldağ,
1800 m, 23.07.1991, 4 ♀♀, leg. H. Özbek, det. H. Özbek;
Cevizli, 15.07.1982, ♂, leg. H. Özbek, det. K. Warncke;
26.07.1983, 3 ♀♀, ♂, leg. H. Özbek, det. B. Tkalcu and
H. Özbek. Erzurum: Gölet, 1950 m, 25.06.2003, ♀, leg. S.
Çoruh, det. H. Özbek; Güngörmez, 2400 m, 05.08.1990,
7 ♂♂, leg. H. Özbek, det. Y. A. Pesenko and H. Özbek;
Karagöbek Mts., 2200 m, 05.07.1989, ♂, leg. H. Özbek, det.
H. Özbek; Köşkköyü, 14.07.1992, ♀, leg H. Özbek, det. H.
Özbek; Şenyurt, 2300 m, 22.08.1991, 10 ♂♂, 6 ♀♀, leg.
H. Özbek and E. Yıldırım, det. H. Özbek, D. Michez, and
W. Thayse; Kargapazarı, Radar road, 2100–2800 m, 20–
28.08.1989, 2 ♀♀, 14 ♂♂, leg. H. Özbek, det. H. Özbek
and Y. A. Pesenko; 11.08.1990, ♀, leg. H. Özbek, det. H.
Özbek; 24.08.1988, 15♀♀, leg. H. Özbek, det. H. Özbek;
Umudum plateau, 2100 m, 14.07.1990, 12 ♀♀, 10 ♂♂,
leg. H. Özbek, det. H. Özbek; Umudum plateau, 2800 m,
08.08.1991, 11 ♀♀, 8 ♂♂, leg. H. Özbek, det. H. Özbek, D.
Michez and W. Thayse; Uzunahmet, 03.08.1981, ♀, leg. H.
Özbek, det. H. Özbek; Aşkale, 07.07.1980, 4 ♀♀, leg. H.
Özışık, det. H. Özbek; Çat, 20.07.1989, ♀, leg. H. Özbek,
det. H. Özbek; Ilıca, Konaklı, 2200–2400 m, 22.07.2000, 5
♀♀, 20 ♂♂, leg. H. Özbek, E. Yıldırım, Ö. Çalmaşur, det.
H. Özbek; Narman, Kireçli Mt., 12.07.2000 m, ♂, leg. C.
Güçlü, det. H. Özbek; Şehitler, 1600 m, 05.08.1997, 2♂♂,
leg. E. Kılıç, det. H. Özbek; Olur, Oğuzkent, 22.08.1992, ♀,
leg. İ. Aslan, det. H. Özbek; Süngübayır, 20.08.1992, ♀, leg.
İ. Arslan, det. H. Özbek; Köprüköy, Örentaş, 04.07.2004,
leg. H. Özbek, det. H. Özbek; Süngübayır, 20.08.1992, ♀,
leg. İ. Aslan, det. H. Özbek; Oltu, Başaklı, 13.08.1977, ♀,
leg. H. Özbek, det. H. Özbek; 01.09.1978, 7 ♀♀, leg. H.
Özbek, det. H. Özbek; Pasinler, Pelitli, 2200 m, 14.08.1996,
5 ♂♂, 2 ♀♀, leg. E. Yıldırım, det. H. Özbek; Rabat, 2400 m,
11.08.1996, 16 ♀♀, 46 ♂♂, leg. E. Yıldırım, det. H. Özbek;
15.08.1995, 4 ♀♀, leg. H. Özbek, det. H. Özbek; Şenkaya,
Soğanlı Pass, 2450 m, 04.08.2000, 2 ♀♀, leg. S. Çoruh,
C. Güçlü, det. H. Özbek; Turnalı, 1750 m, 25.07.1996, ♂,
leg. E. Yıldırım, det. D. Michez and W. Thayse; Turnalı,
06.08.1988, ♂, leg. E. Yıldırım, det. Y. A. Pesenko; Turnalı,
25.08.1991, 3 ♀♀, leg. E. Yıldırım, det. H. Özbek; Turnalı,
2000 m, 01.09.1988 4 ♀♀, ♂, leg. E. Yıldırım, det. H.
Özbek and Y. A. Pesenko. Kars: Sarıkamış, 26.08.1991, 2
♀♀, leg. H. Özbek, det. H. Özbek.
Remark: Radchenko and Pesenko (1989) preferred
using D. thoracica Baer, 1853 for this species, although
none of the specialists accepted changing D. argentata
(Michez et al., 2004a) except for Celary (2002). It is a
widespread species and is very abundant, particularly
ÖZBEK / Turk J Zool
in East Anatolia. After D. hirtipes it is the second most
widespread species of the genus Dasypoda both in Turkey
(Figure 1) and the world.
Plants associated: Daucus carota, Eryngium billardierei,
Cichorium intybus, Echinops galaticus, E. orientalis,
Onopordum acanthium, O. tauricum, Tragopogon aureus,
Cephalaria gigantea, C. lycica, C. procera, Euphorbia
orientalis, Dipsacus laciniatus, Scabiosa caucasica, Melilotus
officinalis, and Salvia aethiopis. It is remarkable to note
that D. argentata shows high preference to the Cephalaria
species; in particular, C. procera is the most frequently
visited plant.
3.1.7. Dasypoda (Megadasypoda) braccata Eversmann,
1852
Distribution: Albania, Austria, Bulgaria, Greece, Caucasus,
Italy, Macedonia, Serbia, European Russia, Hungary,
Poland, Ukraine, Kazakhstan, and Turkey (Warncke, 1973;
Radchenko, 1987; Michez et al., 2004a, 2004b). In Turkey:
Konya, Sivas (Warncke, 1973); Ankara, Kahramanmaraş,
Konya, Niğde, Malatya, Elazığ, Erzincan, Kars, Mardin,
Sivas (Michez et al., 2004a).
Material:
Bayburt:
Demirözü,
31.07.1992,
Bayrampaşa, 31.07.1992, ♀, leg. H. Bostan, D. Michez, and
W. Thayse; Demirözü, 30.07.1992, ♂, leg. H. Bostan, det.
D. Michez and W. Thayse; Kopdağı, 10.08.1991, ♀, leg. H.
Bostan, det. H. Özbek; Yaylapınar, 26.07.1992, ♂, leg. H.
Bostan, det. H. Özbek. Erzincan: Cevizli, 26.07.1983, 2
♀♀, leg. H. Özbek, det. H. Özbek. Erzurum: DSİ Gölet,
25.06.2003, ♀, leg. S. Çoruh, det. H. Özbek; Güngörmez,
2300 m, 05.08.1990, 7 ♀♀, leg. H. Özbek, det. H. Özbek;
Karagöbek, 28.07.1990, ♀, leg. E. Yıldırım, det. H. Özbek;
Kargapazarı, Radar road, 2800 m, 24.08.1988, 12 ♀♀,
leg. H. Özbek, det. H. Özbek; Konaklı, 2000–2400 m,
22.07.2000, ♀, leg. E. Yıldırım, det. H. Özbek; Şenyurt,
2300 m, 22.08.1991, 5 ♀♀, leg. H. Özbek and E. Yıldırım,
det. H. Özbek; Umudum plateau, 2800 m, 08.08.1991,
♂, leg. H. Özbek, det. D. Michez and W. Thayse; Aşkale,
07.07.1980, 5 ♀♀, leg. H. Özışık, det. H. Özbek; Kopdağı,
2200–2400 m, 13.08.1997, 3 ♀♀, leg. Ö. Çalmaşur, det. H.
Özbek; Oltu, Başaklı, 13.08.1977, ♀, 01.09.1978, 4 ♀♀, leg.
H. Özbek, det. H. Özbek; Sütkans, 1500 m 25.06.1996, ♀,
leg. E. Yıldırım, det. D. Michez and W. Thayse; Pasinler,
Rabat, 15.08.1995, ♂, leg. E. Kılıç, det. D. Michez and W.
Thayse; Rabat, 2200 m, 15.08.1995, 2 ♂♂, leg. İ. Aslan, det.
D. Michez and W. Thayse; Oltu, Başaklı-Tutmaç border,
1700–2000 m, 02.07.2000, 13 ♀♀, leg. H. Özbek, det. H.
Özbek; Şenkaya, Turnalı, 25.07.1996, ♂, leg. E. Yıldırım,
det. D. Michez and W. Thayse; Turnalı, 28.07.1993, ♂, leg.
E. Yıldırım, det. H. Özbek; 2000 m, 02.09.1988, 5 ♀♀, leg.
E. Yıldırım, det. H. Özbek; Tortum, Doruklu plateau, 2000
m, Uzundere, Dikyar, 02.07.1994, ♀, leg. İ. Aslan, det. H.
Özbek. Hakkari: Kaval Valley, 1300 m, 20.07.1982, ♂, ♀,
leg. W. Deprins, det. D. Michez and W. Thayse.
Remark: Warncke (1973) described the subspecies D.
braccata anatolica from Konya and Sivas.
Plants associated: Achillea millefolium, Ammi
visnaga, Daucus carota, Eryngium billardierei, Cephalaria
gigantean, C. lycica, C. procera, Scabiosa argentata, S.
caucasica, Carduus acanthoides, C. nutans, Cichorium
intybus, Cirsium arvense, Echinops galaticus, Onopordum
acanthium, O. illyricum. It is particularly associated with
the genus Cephalaria (Dipsacaceae).
3.1.8. Dasypoda (Megadasypoda) frieseana Schletterer,
1890
Distribution: Greece, Macedonia, and Turkey (Warncke,
1973). In Turkey: İçel (Warncke, 1973).
Material: Bayburt: Demirözü, 12.07.1992, 2 ♂♂,
leg. H. Özbek, det. H. Özbek; 30.07.1992, 2 ♀♀, 6
♂♂ leg. H. Özbek, det. D. Michez, W. Thayse and H.
Özbek; 30.07.1992, ♀, ♂, leg. H. Bostan, det. H. Özbek;
31.07.1992, ♀, 4 ♂♂, leg. H. Bostan, det. H. Özbek; 11 ♂♂,
3 ♀♀, leg. H. Özbek, det. D. Michez, W. Thayse and H.
Özbek; 14.08.1992, 3 ♂♂, leg. H. Bostan, det. H. Özbek;
Bayrampaşa, 31.07.1992, ♀, 2 ♂♂, leg. H. Bostan, det. H.
Özbek.
Remark: Warncke (1973) noted that D. friesana was
previously known from Turkey (İçel). However, Baker
(2002) and Michez et al. (2004a) gave the Balkan Peninsula
(Akarnanien) as the only distribution area of this species,
and in the distribution map that was prepared by Michez
et al. (2004a), only Greece and Macedonia were marked.
Recording this species from northeastern Anatolia is
very important under these circumstances. D. frieseana
is recorded in Turkey as well as from the Asian continent
for the first time with the present study and Bayburt is the
easternmost record of this species (Figure 1).
Plants associated: Arctium lappa, Cephalaria gigantean,
C. procera, and Dipsacus laciniatus.
3.1.9. Dasypoda (Megadasypoda) longigena Schletterer,
1890
Distribution: Dasypoda longigena was described from
Anatolia (Kleinasien) by Schletterer (1890). Later it was
recorded from Erzurum (Warncke, 1973; Michez and
Patiny, 2002; Michez et al., 2004a). With the present
knowledge it can be seen that D. longigena is endemic to
Anatolia.
Remark: Dasypoda longigena is one of the earliest bee
species described from Anatolia. It was virtually extinct in
Anatolia and had not been recorded for over 80 years, the
first record after its description being in 1973 by Warncke
(1973) and only in Erzurum. It is remarkable that in spite
of intensive collections in Erzurum and neighboring
provinces since the 1970s it has not been encountered; it is
probably extinct or on the verge of extinction.
449
ÖZBEK / Turk J Zool
3.1.10. Dasypoda (Megadasypoda) patinyi Michez, 2002
Distribution: Syria, Turkey (Michez, 2002; Michez et al.,
2004a). So far it has been recorded in Turkey only from
Şanlıurfa (Michez et al., 2004a).
Remark: Michez (2002) described D. patinyi from
Suwaydan (Syria), which is not far from Şanlıurfa. Dasypoda
patinyi is represented by only one female (Şanlıurfa,
37°08′N, 38°46′E, 01.06.1968, leg. J. Gusenleitner) in
Turkey.
3.1.11. Dasypoda (Megadasypoda) spinigera Kohl, 1905
Distribution: Caucasus, Armenia, Iran, Bulgaria, Greece,
Macedonia, Italy, Romania, Ukraine, Turkey (Warncke,
1973; Michez et al., 2004a); former Russia (Radchenko and
Pesenko, 1989). In Turkey: Kayseri (type locality), Aksaray,
Ankara, Aydın, Bitlis, Çanakkale, İçel, Konya, Nevşehir,
Niğde, Sivas, Van, Tekirdağ (Warncke, 1973; Baker, 2002;
Michez et al., 2004a).
Material: Erzincan: Ilıç, 15.07.1994, ♀, leg. H. Özbek,
det. D. Michez and W. Thayse. Erzurum: Köprüköy,
19.07.1990, ♂, leg. Ş. Güçlü, det. H. Özbek. Konya: Selçuk
University Campus, 14.07.1997, 6 ♀♀, 5 ♂♂, leg. F. Aydın,
det. D. Michez, W. Thayse, and H. Özbek. Van: Edremit,
19.07.1978, ♀, leg. H. Özbek, det. B. Tkalcu.
Plants associated: Cephalaria gigantea, C. procera,
Scabiosa argentata, S. caucasica, and Echinops galaticus.
3.1.12. Dasypoda (Megadasypoda) aurata Rudow, 1881
Synonyms: Dasypoda spectabilis Rudow, 1881; D. mixta
Radoszkowski, 1887; D. thomsoni Schletterer, 1890; D.
suripes (Christ, 1791).
Distribution: Bohemia, Sweden, Denmark, Germany,
Austria, Italy, Poland, Moravia, Slovakia, Hungary, Croatia,
Macedonia, Albania, Romania, Bulgaria, Lithuania,
Greece, Belarus, Ukraine, Armenia, Cyprus, European
Russia, Turkey (Warncke, 1973; Michez et al., 2004a). In
Turkey: Aydın, Hatay, İçel, Muğla (Michez et al., 2004a);
Erzurum as D. mixta Radoszkowski, 1887 (Baker, 2002;
Michez et al., 2004a) (Figure 1).
3.1.13. Dasypoda (Megadasypoda) toroki Michez, 2004
Distribution: Israel, Syria (Michez et al., 2004b).
Material: Antalya: Kumluca, 08.05.1998, 3 ♀♀, leg. G.
Tozlu, det. H. Özbek. Şanlıurfa: Karaköprü, 16.06.1992, ♂,
leg. R. Hayat, det. D. Michez and W. Thayse.
Remark: D. toroki was recently described from Israel
(Michez et al., 2004b). In the present study it has been
recorded from Antalya and Şanlıurfa. Antalya is the
westernmost distribution record of this species (Figure 1).
It is a new record for the Turkish fauna.
Plants associated: Carduus sp.
3.1.14. Dasypoda (Megadasypoda) visnaga (Rossi, 1790)
Distribution: Mediterranean countries from the Atlantic
coast of Portugal to the Aegean coast of Turkey including
North Africa (Algeria, Morocco, and Tunisia). In Turkey:
Antalya (Michez, 2012a).
450
Remark: Although D. visnaga has a very large
distribution range throughout the Mediterranean
countries, including North Africa, more recently it has
been recorded solely from Antalya in Turkey (Figure 1).
3.2. Genus Macropis Panzer, 1809
Macropis is the most distinctive genus of Melittinae; the
front wing has 2 submarginal cells, the face of the male
is yellow, and there is a well-developed pygidial plate in
both sexes (Michener, 1981). Oil-collecting bees are found
worldwide and always in association with particular oilproducing flowers (Bassin et al., 2011). The oil-collecting
and transporting structure of Macropis features short,
dense, velvety hairs on the small segments of all the
tarsi for collecting the oil and the broad hind tibia and
basitarsus with fine, dense, plumose scopal hairs for oil
transport (Michener, 2007). All females of the Macropis
species collect oil and pollen from flowers of Lysimachia
(Primulaceae) as larval food (Rozen and Jacobson,
1980; Cane et al., 1983; Michez and Patiny, 2005). Since
Lysimachia produces no nectar, adults of Macropis visit
other flowers for their own energy needs (Pekkarinen et
al., 2003; Michez and Patiny, 2005).
Macropis includes 16 species in three subgenera:
Paramacropis Popov and Guiglia, 1936; Sinomacropis
Michener, 1981; and Macropis Panzer s. str., occurring
in the eastern Palearctic, in China, and in the Holarctic
region including Turkey, respectively (Michez and Patiny,
2005).
3.2.1. Macropis (Macropis) frivaldszkyi Mocsary, 1878
Distribution: Italy, Bulgaria, Kazakhstan, Macedonia,
Romania, Greece, Croatia, Hungary, Serbia, and European
Russia. In Turkey: Erzurum (Warncke, 1973); Ankara,
Konya, Hakkari, Van (Michez and Patiny, 2005); Sinop
(Bassin et al., 2011) (Figure 2).
Material: Erzurum: Atatürk University Campus,
12.07.1970, ♀, leg. H. Özbek, det. K. Warncke.
Plants associated: Lysimachia vulgaris L. (Primulaceae).
3.2.2. Macropis (Macropis) fulvipes (Fabricius, 1805)
Distribution: Europe (except in the United Kingdom,
Sweden, and Norway), Georgia, Russia, Algeria (Warncke,
1973; Michez and Patiny, 2005; Bassin et al., 2011). In
Turkey: Çankırı, Kastamonu, Artvin, Ardahan, and Rize
(Michez and Patiny, 2005); Kastamonu and Sinop (Bassin
et al., 2011) (Figure 2).
Plants associated: Pekkarinen et al. (2003) noted that
Macropis bees are apparently all oligolectic on Lysimachia;
females collect pollen and oil from Lysimachia, whereas
nectar is usually collected from a wide variety of host
plants. Michez et al. (2008) emphasized that, in the
Palearctic region, Macropis bees are specialized on the
subgenus Lysimachia.
ÖZBEK / Turk J Zool
Macropis frivaldszkyi
Macropis fulvipes
Melitta nigricans
M. bicollaris
M. budensis
M. dimidiata
M. haemorrhoidalis
M. melanura
M. rasmonti
M. singular
M. leporina
Mersin
Figure 2. Distribution of the genera Macropis and Melitta in Turkey.
3.3. Genus Melitta Kirby, 1802
The Melitta species superficially resemble the species of
Andrena (Andrenidae), although their scopae are limited
to the hind tibia and basitarsus (no flocculus). Like other
Melittidae, they have a short tongue with all segments of
the labial palpus similar to one another, one subantennal
suture, and no facial foveae. There are 3 submarginal
cells (the 3rd is longer than the 2nd) in the forewing, no
yellow markings and mainly brownish to blackish pilosity,
widened apical tarsi, lateral tubercles on the labrum, apical
projection on the posterior basitarsus, and volsella with
elongated digitus. In the male, the seventh sternum has a
large disk and weakly developed lateral process (Michener,
1981; Michez et al., 2009).
The genus Melitta occurs in the Palearctic, in southern
and eastern Africa, and in the Nearctic regions (Warncke,
1973; Michener, 1981; Eardley and Kuhlmann, 2006). It
is most diversified in the Palearctic region (Michener,
2007). It is rich in species, although morphologically
monotonous compared to other melittid bees (Michez et
al., 2009).
Warncke (1973) recognized 3 subgenera of Melitta
(Cilissa, Melitta s. str., and Pseudocilissa) based on the
revision of West Palearctic species. Michez and Eardley
(2007) reviewed the literature and examined many
specimens from various collections (including the type
material); they produced the first monographic revision
of the genus Melitta and noted that there are currently
48 valid species. However, very recently, Dellicour et al.
(2014) described 2 more Melitta species, and so it rose to
50 species.
3.3.1. Melitta (Melitta) leporina (Panzer, 1799)
Synonyms: Apis leporina Panzer, 1799; Andrena fortipes
Imhoff, 1832; Cilissa ruthenica Radoszkowski, 1891;
Melitta centaureae Torka, 1922
Distribution: Palearctic. Widespread in Europe
including southern Sweden and Finland. In Asia, known
from Anatolia and Iran, Kyrgyzstan to Mongolia (Warncke,
1973; Michez and Eardley, 2007). In Turkey: Erzurum
(Warncke, 1973; Özbek, 1976, 1979, 1980).
Material: Afyon: Sultandağı, 25.06.1995, ♂, leg. H.
Özbek, det. H. Özbek. Aksaray: Center, 27.06.1975, 4 ♀♀,
2 ♂♂, leg. H. Özbek, det. H. Özbek. Amasya: Merzifon,
23.07.1998, ♂, leg. H. Özbek, det. H. Özbek. Artvin:
Yusufeli, 1350 m, 22.07.2003, 3 ♀♀, leg M. Kesdek, det.
H. Özbek; Cinnar, 15.07.1992, 2 ♂♂, leg. E. Yıldırım,
det. H. Özbek. Bilecik: Center, 21.06.1978, 3 ♂♂, leg. H.
Özbek, det. H. Özbek. Bursa: Uludağ, 20.06.1978, ♀, 3
♂♂, leg. H. Özbek, det. H. Özbek. Erzincan: Çağlayan,
39.56670_39.65000, 06.07.2001, ♂, leg. J. G. Rozen and
H. Özbek, det. D. Michez. Erzurum: Atatürk University
Campus, 39.90250_41.23500, 2000 m, 07.06.2007, ♂, leg.
J. S. Ascher, H. Özbek and J. G. Rozen, det. J. S. Ascher;
28.06.1978, ♂, leg. H. Özbek, det. H. Özbek; 28.06.2001, ♂,
leg. J. G. Rozen and H. Özbek, det. D. Michez; 30.06.2007,
♀, leg. J. S. Ascher, H. Özbek, J. G. Rozen, det. J. S. Ascher;
17.07.2010, 4 ♀♀, leg. J. S. Ascher, H. Özbek, det. J. S.
Ascher; 21.07.2010, 2 ♀♀, ♂; 29.07.2010, ♂, leg. J. S.
Ascher, H. Özbek, J. G. Rozen, det. J. S. Ascher; 10.07.2001,
♀, leg. H. Özbek, J. G. Rozen, det. D. Michez; 17.07.2003,
2 ♀♀, ♂, leg. J. G. Rozen and H. Özbek, det. D. Michez;
18.07.2003, 2 ♀♀, leg. J. G. Rozen and H. Özbek, det. D.
Michez (in AMNH); 19.07.1987, ♂, leg. R. Hayat, det. H.
Özbek; 21.07.2003, 3 ♀♀, leg. J. G. Rozen and H. Özbek,
det. D. Michez; 12.07.1970, 2 ♀♀, leg. H. Özbek, det. K.
Warncke; 12–15.07.1971, 12 ♀♀, leg. H. Özbek, det. K.
Warncke; 20–26.07.1971, 10 ♀♀, leg. H. Özbek, det. K.
Warncke and H. Özbek; 17–22.07.1974, 5♀♀, leg. H.
451
ÖZBEK / Turk J Zool
Özbek, det. K. Warncke; 29.07.1974, 4 ♀♀, leg. H. Özbek,
det. K. Warncke; 20–29.07.2003, 10 ♀♀, leg. H. Özbek,
det. H. Özbek; İspir, 15.07.1991, ♀, leg. H. Özbek, det. K.
Warncke; İspir, Kan, 20.07.1970, ♀, leg. H. Özbek, det. K.
Warncke; Oltu, 4 km WSW of Oltu, 40.53305_41.98527,
1500 m, 03.07.2001, 2 ♂♂, leg. J. G. Rozen and H. Özbek,
det. D. Michez; 30.06.1980, ♀, leg. H. Özbek, det. K.
Warncke; 04.07.1974, 4 ♀♀, leg. H. Özbek, det. K. Warncke
and H. Özbek; 08–11.07.1974, 36 ♂♂, leg. H. Özbek, det.
H. Özbek; 9.07.1970, 2 ♀♀, 2 ♂♂ leg. H. Özbek, det. K.
Warncke and H. Özbek; 12–20.07.1970, 8 ♀♀, 4 ♂♂ leg.
H. Özbek, det. K. Warncke; 14–11–17.07.1974, 25 ♂♂, leg.
H. Özbek, det. H. Özbek; 18.07.1970, 2 ♂♂, leg. H. Özbek,
det. H. Özbek; 18.07.1972, 2 ♂♂, leg. H. Özbek, det. H.
Özbek; 20.07.1971, 7 ♂♂, leg. H. Özbek, det. H. Özbek;
23.07.1974, leg. H. Özbek, det. H. Özbek; 24.07.1970, ♀,
leg. H. Özbek, det. K. Warncke; 26–28.07.1972, 9 ♀♀, leg.
H. Özbek, det. H. Özbek; 26.07.1966, ♀, leg. H. Özbek,
det. K. Warncke; 29.07.1970–74, 5 ♂♂, leg. H. Özbek, det.
K. Warncke; Kandilli, 29.06.1970, ♂, leg. H. Özbek, det.
H. Özbek; Olur, Coşkunlar, 12.07.1991, ♀, leg. E. Yıldırım,
det. H. Özbek; Oltu, 20.07.1970, 4 ♂♂, leg. H. Özbek, det.
H. Özbek; 19.08.1987, ♀, ♂, leg. H. Özbek, det. H. Özbek;
Tortum, Pehlivanlı, 12.07.1992, ♂, leg. E. Yıldırım, det. D.
Michez. Eskişehir: Sivrihisar, 13.08.1990, 2 ♂♂, leg. H.
Özbek, det. H. Özbek. Iğdır: Aralık, 02.08.1997, 4 ♂♂,
leg. B. Gül, det. H. Özbek. İzmir: Bornova, 08.07.1979,
♀, ♂, leg. H. Özbek, det. H. Özbek. Kayseri: Develi,
27.06.1975, 5 ♀♀, 3 ♂♂, leg. H. Özbek, det. H. Özbek.
Kütahya: Tavşanlı, 28.06.1979, ♂, leg. H. Özbek, det. H.
Özbek. Manisa: Salihli, 05.06.1989, ♀, leg. H. Özbek, det.
H. Özbek. Karaman: Center, 02.07.1975, 2 ♀♀, 4 ♂♂, leg.
M. Kesdek, det. H. Özbek. Konya: Güneysınır, Gürağaç,
1017 m, 23.06.1999, ♀, leg. M. Kesdek, det. H. Özbek.
Nevşehir: Ürgüp, 28.06.1975, 2 ♀♀, 3 ♂♂, leg. H. Özbek,
det. H. Özbek. Niğde: Çamardı, 29.06.1975, 3 ♀♀, 2 ♂♂,
leg. H. Özbek, det. H. Özbek. Samsun: Kavak, 26.07.1999,
2 ♀♀, ♂, leg. H. Özbek, det. H. Özbek. Sinop: Gerze,
13.07.1998, ♀, 3 ♂♂, leg. H. Özbek, det. H. Özbek. Sivas:
Hafik, 17.06.2001, ♂, leg. H. Özbek, det. H. Özbek. Van:
Muradiye, 38.99000_43.76000, 1750 m, 16.07.1988, ♀,
leg. Schmid-Egger, det. K. Warncke (in AMNH). Yozgat:
Akdağmadeni, 08.07.2003, ♂, leg. H. Özbek, det. H.
Özbek.
Remark: M. leporina is the most widespread species for
the genus; it occurs from England to Vladivostok and from
Finland to Spain (Michez, 2010). It is also one of the most
widespread and abundant bee species occurring in Turkey
(Figure 2).
Plants associated: Melitta leporina visits alfalfa
(Medicago sativa L.), sainfoin (Onobrychis viciifolia Scop.),
red clover (Trifolium pratense L.), and white clover (T.
repens L.), which are important fodder crops growing in
452
Turkey. Wild plants: Fabaceae: Astragalus aduncus Willd.,
A. aureus Willd., A. baibutensis Bunge, A. bicolor Lam.,
A. christianus L., A. galegiformis L., A. lagurus Willd.,
A. longifolius Lam., A. onobrychis L., A. ponticus Pall.,
Coronilla orientalis Mill., C. varia L., Glycyrrhiza glabra
L., Hedysarum elegans Boiss. & A.Huet, H. nitidum Willd.,
Lathyrus spp., Lotus corniculatus L., Medicago falcata L.,
M. lupulina L., M. papillosa Boiss., Melilotus alba Desr., M.
officinalis (L.) Desr., Onobrychis armena Boiss. & A.Huet.,
O. cana Boiss., Trifolium ambiguum M.Bieb., T. montanum
L., Vicia cracca L., and V. villosa Roth.
Celary (2006) described the nesting ecology of M.
leporina. The female builds its nest in hard, clayed soil.
They are not gregarious and the entrance to each nest is
concealed by vegetation.
3.3.2. Melitta (Melitta) nigricans Alfken, 1905
Distribution: Widespread in Europe from Portugal
to Poland and Orenburg (Russia). Absent in England
and Sweden, one record from Morocco and Argentina
(Warncke, 1973; Michez and Eardley, 2007). In Turkey:
Balıkesir (Warncke, 1973).
Remark: Although M. nigricans is a common species
distributed throughout Europe, except for FennoScandinavia, it is very rare in Turkey, known only in
Balıkesir (Figure 2).
Plants associated: Michez et al. (2008) gave the family
Lythraceae as the host plants of M. nigricans.
3.3.3. Melitta (Cilissa) bicollaris Warncke, 1973
Distribution: Warncke (1973) described the female of M.
bicollaris from Erzurum. Later, Michez and Eardley (2007)
described the male based on a pair of specimens collected
from Erzurum. So far it has been known only from Artvin,
Erzincan, Erzurum, and Kars provinces (Figure 2). With
the present data it is endemic to East Anatolia.
Material: Erzurum: Atatürk University Campus,
09.07.1976, 2 ♀♀, leg. H. Özbek, det. H. Özbek; 15.07.1972,
2 ♀♀, leg. H. Özbek, det. K. Warncke; 13.07.1972, 5 ♀♀,
leg. H. Özbek, det. K. Warncke and H. Özbek; 18.07.1972,
5 ♀♀, 11 ♂♂, leg. H. Özbek, det. H. Özbek; 28.07.1972,
10 ♀♀, leg. H. Özbek, det. H. Özbek (all samples on O.
viciifolia); 2000 m, 39.90250_41.23500, 02.07.2002, ♂,
leg. J. G. Rozen and H. Özbek, det. D. Michez; Umudum
plateau, 2200 m, 14.07.1990, 12 ♀♀, leg. H. Özbek and Ş.
Güçlü, det. H. Özbek; Tortum, Pehlivanlı, 12-29.07.1992,
2 ♀♀, leg. E. Yıldırım, det. D. Michez; Şenkaya, Turnalı,
06.07.1990, 2 ♂♂, leg. E. Yıldırım, det. H. Özbek.
Plants associated: Astragalus aduncus, A. bicolor, A.
gummifer, A. lagurus, A. ponticus, Hedysarum elegans,
Lotus corniculatus, Medicago papillosa, M. sativa, Melilotus
alba, M. officinalis, Onobrychis cana, O. cf. stenostachya,
O. transcaucasica, O. viciifolia, Medicago sativa, Trifolium
montanum, T. repens, and Vicia cracca.
ÖZBEK / Turk J Zool
3.3.4. Melitta (Cilissa) budensis (Mocsary, 1878)
Distribution: Hungary, Russia, Ukraine, Armenia,
Kazakhstan, Turkey (Warncke, 1973; Michez and Eardley,
2007). In Turkey: Bolu (Warncke, 1973); Bitlis, Bolu,
Erzurum, Hakkari (Michez and Eardley, 2007) (Figure 2).
Plants associated: Vicia cracca (Michez and Eardley,
2007).
3.3.5. Melitta (Cilissa) dimidiata Morawitz, 1876
Synonyms: Cilissa robusta Radoszkowski 1876, Cilissa
dimidiata hungaria Mocsary 1883, Pseudocilissa robusta,
Radoszkowski (1891).
Distribution: South and East Europe as far as Central
Asia (Warncke, 1973; Michez and Eardley, 2007). Some
isolated populations in England (Baker, 1965). In Turkey:
Ankara, Bitlis, Erzurum, İçel, Konya (Warncke, 1973;
Özbek, 1979, 2011) (Figure 2).
Material: Erzurum: Atatürk University Campus,
02.06.1974, ♀, leg. H. Özbek, det. H. Özbek; 04.06.1970,
2 ♀♀, ♂, leg. H. Özbek, det. H. Özbek and K. Warncke;
08.22.06.1970, 55 ♂♂, 16 ♀♀, leg. H. Özbek, det. K.
Warncke and H. Özbek; 09.06.1970, ♂, leg. H. Özbek, det.
K. Warncke; 24.06.1970, 2 ♀♀, leg. H. Özbek, det. H. Özbek;
20.06.1971, 6 ♀♀, leg. H. Özbek, det. H. Özbek; 30.06.197,
2 ♀♀, leg. H. Özbek, det. H. Özbek; 02–20.07.1970, 3 ♀♀,
♂, leg. H. Özbek, det. H. Özbek; 06–26.07.1971, 48 ♂♂,
24 ♀♀, leg. H. Özbek, det. H. Özbek and K. Warncke; 02–
29.06.1974, 39 ♂♂, ♀, leg. H. Özbek, det. H. Özbek and B.
Tkalcu; 01.07.1972, 3 ♂♂, leg. H. Özbek, det. H. Özbek;
06.07.1976, ♂, leg. H. Özbek, det. H. Özbek; 12.07.1970,
♀, leg. H. Özbek, det. H. Özbek; 16.07.1966, 2 ♀♀, leg. H.
Özbek, det. H. Özbek; 11.07.1967, ♂, leg. H. Özbek, det. B.
Tkalcu; 17.07.1972, 3 ♀♀, leg. H. Özbek, det. H. Özbek and
B. Tkalcu; 20.07.1971, 3♀♀, leg. H. Özbek, det. B. Tkalcu;
26.07.1974, ♀, leg. H. Özbek, det. H. Özbek; 10.06.2000,
3 ♂♂, leg. M. Kesdek, det. H. Özbek; 03.07.1992, 5 ♀♀,
♂, leg. E. Yıldırım, det. H. Özbek; Umudum plateau,
2200 m, 14.07.1990, 2100 m, 2 ♂♂, leg. Ş. Güçlü, det. D.
Michez; 08.08.1991, ♂, leg. H. Özbek, det. D. Michez;
2200 m, 14.07.1991, 2 ♂♂, leg. Ş. Güçlü and E. Yıldırım,
det. D. Michez; İspir, 15.06.1971, ♂, leg. H. Özbek, det.
H. Özbek; Oltu, center, 25.06.1971, 2♀♀, leg. H. Özbek,
det. H. Özbek; 14.07.1972, 2 ♀♀, ♂, leg. H. Özbek, det.
H. Özbek; Başaklı, 15.07.1989, ♂, leg. H. Özbek, det. D.
Michez; Şenkaya, Turnalı, 01.07.1990, ♂, leg. E. Yıldırım,
det. H. Özbek; Tekman, Yayla, 19.07.2005, 2450 m, ♂, C.
Güçlü, det. H. Özbek, Tortum, 14.07.1972, 3 ♀♀, ♂, leg.
H. Özbek, det. H. Özbek; Uzundere, 28.06.1992, leg. M.
Atamanalp, det. H. Özbek.
Plants associated: Medicago sativa, O. viciifolia, T.
pratense, T. repens. The present observations in the field
and previous studies (Özbek, 1979, 2011) reveal that M.
dimidiata is one of the most important pollinators of
sainfoin (O. viciifolia), which is a valuable fodder crop
growing mainly in the East and Central Anatolian regions
of Turkey. Wild plants: Astragalus aduncus, A. bicolor,
A. cylindraceus, A. gummifer, A. lagurus, A. ponticus,
Hedysarum elegans, H. nitidum, Lotus corniculatus,
Medicago papillosa, M. sativa, Melilotus alba, M. officinalis,
Onobrychis altissima, O. arenaria, O. araxina, O. cana,
O. cornuta, O. transcaucasica, Ononis spinosa, Trifolium
ambiguum, T. montanum, Vicia alpestris, V. cracca.
3.3.6. Melitta (Cilissa) haemorrhoidalis (Fabricius, 1775)
Synonyms:
Andrena
haemorrhoidalis
Fabricius,
1775; Cilissa haemorrhoidalis (Fabricius, 1775); Apis
dichroa Gmelin, 1790; M. chrysura Kirby, 1802; M.
haemorrhoidalis nigra (Friese, 1885); M. haemorrhoidalis
frisica Zimmermann, 1935; M. haemorrhoidalis flavidorsis
Ruszkowski, 1988
Distribution: Armenia, Austria, Belgium, Bohemia,
Bulgaria, Czech Republic, Denmark, Estonia, European
Russia, France, Finland, Germany, Georgia, Greece,
Hungary, Italy, Latvia, Lithuania, Luxembourg, Moravia,
Netherlands, Norway, Poland, Romania, Slovakia,
Slovenia, Spain, Sweden, Switzerland, Ukraine, United
Kingdom (Michez, 2012b). Although M. haemorrhoidalis
is one of the most common and widespread Melittidae
species in Europe, recently it was recorded in Rize
(İkizdere, 40.78333_40.55) by Michez (2012b) (Figure 2).
Plants associated: Michez and Eardley (2007) recorded
M. haemorrhoidalis mainly on Campanula latifolia, C.
persicifolia, C. rapunculoides, C. rotundifolia, and C.
trachelium (Campanulaceae); Malva moschata and M.
sylvestris (Malvaceae).
3.3.7. Melitta (Cilissa) melanura (Nylander, 1852)
Synonyms: Kirbya melanura Nylander, 1852; Cilissa
wankowiczi Radoszkowski, 1891; Melitta wankowiczi
(Radoszkowski, 1891).
Distribution: Austria, Belarus, European Russia,
Greece, Germany, Lithuania, Mongolia, Poland, Romania,
Slovakia, Sweden, Ukraine, Turkey (Michez, 2012b).
Although M. melanura is widespread in the Palearctic
from Austria to Mongolia, in Turkey it has been recorded
only in Kars so far (Figure 2). It is represented only by
one specimen [1♀, Kars, Sarıkamış (40.33°N, 42.58°E),
06.08.1979 as M. wankowiczi (Radoszkowski, 1891)]
(Michez and Eardley, 2007).
3.3.8. Melitta (Cilissa) rasmonti Michez, 2007
Distribution: It was described in Hakkari, and one
specimen was recorded in Erzurum (Palandöken,
15.08.1978, ♀, leg. H. Özbek) (Figure 2). It is a mountain
species; all materials were collected at an elevation of 2300
m or higher.
3.3.9. Melitta (Cilissa) singular Michez, 2012
Distribution: only known from the type locality in Turkey
(Kırşehir, Mucur, 1000 m, ♂, 25.06.1988, leg. K. Guichard)
(Figure 2). It is represented only by one specimen.
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4. Discussion
The examination of samples collected from various parts
of the country and an overview of the literature allow us to
reach the conclusion that the Melittidae of Turkey comprise
25 species in 3 genera: Dasypoda (14 species), Macropis (2
species), and Melitta (9 species). The Melittidae include
approximately 200 species in 14 genera (Michez et al.,
2009, 2012) worldwide. With 25 species, Turkey is home
to more than 12% of the world’s Melittidae. Among these
species, the most common and most frequently observed
species across Turkey are Dasypoda hirtipes, D. argentata,
D. braccata, Melitta leporina, and M. dimidiata. These
species are also widespread outside of Turkey; they have
Palearctic distribution. However, certain species are very
rare and most are known from only 1 or 2 localities and
mainly from 1 locality, including D. friesana (from İçel and
Bayburt), D. longigena (from Erzurum), D. patinyi (from
Şanlıurfa), D. visnaga (from Antalya), Melitta nigricans
(from Balıkesir), M. haemorrhoidalis (from Rize), M.
rasmonti (from Erzurum and Hakkari), M. singular (from
Kırşehir), and M. melanura (from Kars). Interestingly, M.
melanura and D. visnaga are widespread in the Palearctic,
although they have been recorded only from one locality
each. Additionally, M. melanura and M. singular are
represented only by one specimen each. Concerning new
records, D. toroki, recently described from Israel and Syria
(Michez et al., 2004b), was recorded from Antalya and
Şanlıurfa provinces.
Eight species (more than 30% of the Turkish melittids),
5 Dasypoda and 3 Melitta, have type localities from
Anatolia: Dasypoda tubera (İçel), D. warnckei (Kayseri),
D. longigena (Anatolia, locality uncertain), D. pyrotrichia
(Anatolia, locality uncertain), D. spinigera (Kayseri),
Melitta bicollaris (Erzurum), M. rasmonti (Hakkari), and
M. singular (Kırşehir). Of these, D. longigena, D. warnckei,
M. bicollaris, M. singular, and M. rasmonti are endemic
to Anatolia (20% of the total species). Dasypoda spinigera
and D. pyrotrichia are quite widespread both in Turkey and
outside of Turkey. Dasypoda pyrotrichia has been recorded
mostly from the northeastern part of Anatolia. Michez
(2012) pointed out that 3 subspecies of D. pyrotrichia are
recognized in 3 different areas around the Mediterranean
basin: Spain, Greece/Bulgaria, and Anatolia. Dasypoda
longigena and D. friesana were described by Schletterer
(1890) from Turkey and Greece, respectively. No material
has been found since the description of both of these
species in Turkey. However, Warncke (1973) added new
localities for D. friesana from Greece and Macedonia.
Unfortunately, they appear to be either extinct or on the
verge of extinction in Turkey.
In general, when we take a look at the distribution
range of the Melittidae fauna of Turkey, all the
species occurring in Turkey are associated within the
454
Mediterranean, particularly the East Mediterranean basin.
The Mediterranean is a vast, biologically rich region,
comprising Mediterranean ecosystems, arid and semiarid
coastal and inland ecotones, and landscapes of faunistic
and floristic richness set in a geographically diverse mosaic.
The topography is varied, with many mountainous regions
but also with low-lying plains and coastal strips. This could
explain the high diversity of plants and animals as well as
Melittidae bees in Anatolia. I suggest that the speciation
of most of these species occurred in the Mediterranean
basin, including Anatolia, and then dispersed to various
directions. In their dispersal, apart from paleogeographic
and paleoclimatic events, dispersal power (being large in
size with a strong body structure) and the adaptive features
of these species could play an important role.
It is worth emphasizing that the current knowledge
on the distribution and fauna of the family Melittidae in
Turkey is still incomplete. There are approximately 10
species known from only 1 locality each. Furthermore,
the occurrence of several species that are known to occur
in neighboring countries is suspected. In particular, the
occurrence in Turkey of the following species seems likely
or at least possible: Dasypoda litigator Bakker, 2002 and
D. intermedia Michez, 2005 were recently described from
Iran; D. syriensis Michez, 2004 was described from Syria;
and D. cingulata Erichson, 1835 occurs in Mediterranean
countries including Greece. I think that all of these species
theoretically could occur in Turkey, although intensive
collections must be conducted in various places, in
particular at the appropriate time of the year and on the
right host plant.
4.1. Melittidae bees as pollinators of cultivated and wild
plants
4.1.1. Pollinators of crops
Mutualisms between insects and plants are widely
recognized as indispensable components in ecosystems
functioning (Bronstein et al., 2006). Agricultural
pollination is the first indispensable step in a process that
results in the production of fruits, vegetables, nuts, and
seeds. Therefore, agriculture is highly dependent on insect
pollination (McGregor, 1976; O’Toole and Raw, 1991;
Free, 1993; Cunningham et al., 2002; Woodcock, 2012;
Garibaldi et al., 2013). Bees are present in most ecosystems,
where they are primarily responsible for the pollination of
flowering plants, playing a key role in ecosystems dynamics.
It is a well-known fact that more than 35% of global crop
production depends on insect pollination. Most of these
pollinators are wild bees (Pedersen, 1961; Palmer-Jones
et al., 1966; Corbet et al., 1993; Kremen et al., 2002;
Velthuis and Doorn, 2006; Klein et al., 2007; Garibaldi
et al., 2013). Wild bees are of enormous ecological and
economic importance as pollinators of many cultivated
and wild plants and are thought to account for 75% of
ÖZBEK / Turk J Zool
crop pollination requirements (Nabhan and Buchmann,
1997); hence, they are often termed a keystone species
(O’Toole, 1993). Only a dozen bee species are managed
for pollination worldwide (Kremen et al., 2002); in
contrast, hundreds to thousands of the world’s more than
20,000 wild bee species contribute to crop pollination as
unmanaged populations (McGregor, 1976; Corbet et al.,
1991; Free, 1993; Nabhan and Buchmann, 1997). It has
often been observed that alternative bee species are either
equally efficient or more efficient than the most widely
used managed pollinator, Apis mellifera L., in a variety of
crops (alfalfa, blueberry, red clover, cranberry, sunflower,
watermelon, tomato, and many more) (Parker et al., 1987;
Kevan et al., 1990; Woodcock, 2012).
Woodcock (2012) indicated that even plants that are
considered self-fertile can often realize a benefit in the
quantity or quality of production from cross-pollination,
or even from the activity of insects moving pollen around
on the flower that results in more grains being delivered.
Even in self-compatible plants, self-pollination is often an
“emergency mechanism” used by the plant to assure some
reproduction if cross-pollination fails. Seeds and fruits
that develop as a result of self-pollination are often smaller
and of lower quality than those resulting from crosspollination, because the plant will preferentially invest
resources in cross-pollinated offspring and others may
even be aborted, reducing yield.
Like many groups of phytophagous insects (Jaenike,
1990), bees show variation in host-plant use as well as hostplant breadth. Whereas many bee species exhibit floral
specificity, visiting only a restricted number of suitable and
available plant taxa throughout their range (i.e. monolecty
or oligolecty), others display a wider spectrum of pollen
hosts (i.e. mesolecty or polylecty) (Michez et al., 2008). In
general, when we examine the Melittidae species associated
with plant species, Dasypoda has a strong preference for
the family Asteraceae, Melitta has a strong preference for
Fabaceae, and Macropis is monolectic on Primulaceae.
Under these circumstances, as a family, Melittidae display
a polylectic structure.
Among the cultivated plants, sunflower, Helianthus
annuus, and safflower, Carthamus tinctorius, were visited
by D. hirtipes, D. pyriformis, and D. braccata. Particularly,
D. hirtipes has been found to be the most intense and
widespread species and frequently visits these plants.
Different cultivars of sunflowers exhibit different degrees of
self-fertility. The isolated heads usually hold approximately
10%–20% seeds, compared with 70%–90% in exposed
heads (Free, 1993). Observations and experiments on
the field scale confirm the value of bees in pollinating
sunflowers and demonstrate that a shortage of bees may
limit sunflower seed production (Free, 1993). Although
the honey bee (A. mellifera) is usually the most abundant
bee species visiting sunflowers and constitutes the highest
percentage, Çalmaşur and Özbek (1999) reported that
many locally abundant bees contribute to pollination and
may be major pollinators. These researchers found that
of 42 bee species in the 5 families visiting sunflowers in
Erzurum, honey bees accounted for approximately 80%,
with wild bees representing 20%. Free (1993) noted that
although relatively few in number, solitary bees are the
most efficient pollinators of sunflowers. Whereas honey
bees often show little interest in collecting sunflower
pollen, solitary bees often concentrate on doing so.
Safflower is commercially cultivated for vegetable oil
extracted from its seeds. It is now grown as an oil crop
in Turkey and some other countries. Boch (1961) caged
3 plots of safflowers to exclude bees and found that
they produced only approximately half as much seed as
uncaged plots. He found that 90% of the insects visiting
safflowers were honey bees. Various researchers found
that, besides honey bees, different wild bee species visited
safflowers (Free, 1993). Eckert (1962) found that a selffertile cultivar of safflowers set as much seed when caged
to exclude insects as in the open (66: 64% set), although
a cultivar that was a somewhat self-sterile set less when
caged than in the open (41: 59% set). Dasypoda hirtipes
visits safflowers; it could be one of the efficient pollinators
in favorable conditions in Turkey.
Chicory, Cichorium intybus, is a light and refreshing
vegetable used in salads and sandwiches; the flowers are
also edible for salads and they can be frozen in ice cubes
that are then added to iced tea. This plant is very abundant
and widespread in Turkey, but cultivation might be very
restricted. Most of the Dasypoda species, including D.
tubera, D. warnckei, D. argentata, D. braccata, D. friesana,
and D. spinigera, and, in particular, D. hirtipes, visit C.
intybus flowers, from which they mostly collect pollen.
It is evident from the present study that the genus
Melitta is shown to be polylectic, although it exhibits
a clear preference for the family Fabaceae. Among
the Melitta species, M. leporina, M. dimidiata, and M.
bicollaris play an important role in the pollination of
Fabaceae plants, such as alfalfa (M. sativa), sainfoin
(O. viciifolia), red clover (T. pratense), white clover (T.
repens), soybean (Glycine max L.), beans (Phaseolus spp.),
pea (Pisum sativum L.), chickpeas (Cicer arietinum L.),
peanut (Arachis hypogaea L.), and licorice (Glycyrrhiza
glabra L.) (Free, 1993). Pollination has frequently been
the principal limiting factor in the growing of these
plants, particularly alfalfa and red clover, for seed. Field
observations, especially in East and Central Anatolia, and
previous studies (Özbek, 1976, 1979, 2008) revealed that
M. leporina is a valuable pollinator of alfalfa in Turkey.
The flower of alfalfa is peculiar, constructed so that the
staminal column is held under pressure within the keel
455
ÖZBEK / Turk J Zool
by interlocking projections from the keel and wing petals.
When the tension is released, the staminal column snaps
forward against the standard petal, causing the pollen to
be distributed (Pedersen, 1961). This process is known as
“tripping”, and once it has occurred, the staminal column
does not return to the keel. Tripping is accomplished when
the keel is pressed down by the weight of a visiting bee
(Pedersen, 1961; Özbek, 1979; Free, 1993). Individual
alfalfa plants range from being completely self-sterile or
completely self-fertile, although more seeds are formed per
pod from cross-pollination than from self-pollination and
seeds from cross-pollination are larger (Free, 1993). Özbek
(1979) found that in caged pods, setting was 5.48% and
seeds per pod were 1.42, while in exposed conditions they
were 64.49% and 4.34, respectively. Although the relative
importance of honey bees and wild bees differs greatly
in different localities, a high proportion of honey bees
visiting alfalfa flowers fail to pollinate them, particularly
in locations where competing plants are abundant (Free,
1993). I observed that honey bees rarely visit alfalfa flowers
and fail to collect pollen in Erzurum due to the abundance
of other flowering plants such as O. viciifolia, Melilotus
alba, M. officinalis, and T. repens in the area. Likewise,
Özbek (1979) noted that wild bees were important in the
pollination of alfalfa and listed 26 species visiting alfalfa
flowers in Erzurum Province. Later, in his countrywide
study, he found that approximately 150 species of bees in 5
families of 30 genera visited alfalfa flowers in Turkey, which
is located in the area where alfalfa supposedly originated
(Özbek, 2008). Among the Melittidae species, M. leporina
was one of the most efficient pollinators of alfalfa. It is
widespread in Turkey (Figure 2), active from sunrise to
sunset during the flowering period and visiting 14–16
alfalfa flowers per minute. I made a field trip to Central
Anatolia, which is the main alfalfa seed-growing area of
Turkey, to detect the activities and pollinating efficiency
of M. leporina and make demonstrations in the field to the
experts at the Agricultural Ministry. We observed together
that it is very abundant in Central Anatolia and active
throughout the flowering period, efficiently pollinating
alfalfa flowers. The experts tracked the activities of M.
leporina and revealed the importance of this bee in the
pollination of the alfalfa plant. Free (1993) noted that M.
leporina is a particularly good alfalfa pollinator in some
other countries, such as Bulgaria, Denmark, France,
Hungary, Poland, Sweden, and the former USSR.
In contrast to M. leporina, 2 other Melitta species, M.
dimidiata and M. bicollaris, rarely visit alfalfa, especially
in the presence of sainfoin (O. viciifolia) and some other
competing plants such as Melilotus alba, M. officinalis,
Lotus corniculatus and T. repens in the surroundings.
Field observations and my previous studies (Özbek, 1979,
2008) revealed that both of these species show a definite
456
preference for sainfoin pollens and, while collecting the
pollens, pollinate about 90% of the flowers visited. The
population of M. dimidiata is much larger than that of M.
bicollaris, which has a distribution area restricted to the
eastern part of the country. Regarding sainfoin pollinators,
M. dimidiata is the most abundant and widespread wild
bee; it visited 13–15 flowers per minute and foraged at
0600–1800 hours (Özbek, 1979). Among the wild bees it
could be the best pollinator of sainfoin in Turkey. Although
it is slightly self-fertile, it is a cross-pollinated plant.
Clovers (Trifolium spp.) are also important crops with
high value as livestock forage, green manure, and cover
crops and also as honey plants in Turkey. Red clover
(T. pratense), white clover (T. repens), and alsike clover
(T. hybridum) are the major species growing in Turkey
(Açıkgöz, 2001). The flowers of clover follow the typical
legume structure, although the individual tubular flowers
are small, narrow, and grouped together in inflorescences.
The anthers dehisce and release the pollen inside the bud
prior to opening. The weight of a flower visitor exerts
pressure on the standard and wing petals and causes the
anthers and stigma to extend forward and press against the
underside of the head of the visitor (Free, 1993). Following
a visit, in contrast to the alfalfa flower, the sexual structures
return to their original position, allowing the same flower
to deliver pollen repeatedly (Bohart, 1957). The action of
insects is required for a successful seed set. Manipulation
of the flowers causes pollen to be deposited on the ventral
surface of the pollinator bee at the same time that the
stigma is contacted and cross-pollen delivered. Although
the main pollinators of Trifolium spp., especially T.
pratense, are various species of bumble bees and honey
bees (Bohart, 1957; Palmer-Jones et al., 1966; Özbek,
1980; Woodcock, 2012), M. leporina, M. dimidiata, and
M. bicollaris visit T. pratense, T. repens, and T. hybridum.
Field observations revealed that the presence of competing
plants in the vicinity of clover-growing area caused visiting
Melitta species to be less frequent. The main competing
plants are sainfoin (for M. dimidiata and M. bicollaris) and
alfalfa (for M. leporina). My impression is that T. repens is
more attractive for 3 of the Melitta species than the other
Trifolium species.
In conclusion, although the pollination of alfalfa,
sainfoin, and clover species appear to present some
difficulties, M. leporina is one of the most efficient
pollinators of alfalfa, although it is less attracted to sainfoin
and clover species. Melitta dimidiata and M. bicollaris,
and especially the former, are very efficient pollinators
of sainfoin, although they are moderately important for
clover species. Furthermore, alfalfa is less attractive to M.
dimidiata and M. bicollaris.
It is clear that increasing floral abundance and plant
diversity reduced pollinator visitations to the target plants,
ÖZBEK / Turk J Zool
suggesting competitive effects through both the quantity
and diversity of resources. As pollinating agents of these
leguminous plants, in order to obtain more benefits
from these bee species, efforts should be made to avoid
competition with other flower species. Additionally,
Melitta species mostly nest in field margins (Celary, 2006).
Therefore, in seed-growing areas, field margins, field edges
and paths, headlands, fence-lines, and nearby uncultivated
patches of lands are important refuges for them.
4.1.2. Pollinators of wild plants
It is evident from this study and the literature data that, in
addition to cultivated plants, the members of the family
Melittidae visit many wild plant species in the various
plant families and pollinate them. In order to conserve
and maintain biodiversity and to keep ecology in good
conditions, in addition to cultivated plants, wild plant
species also have to be pollinated at the optimum level.
Furthermore, pollinator bees are indirectly responsible
for the persistence of other guilds that depend upon floral
resources, such as herbivores and seed-eaters.
Dasypoda species show a large variation in the hostplant choices. Michez et al. (2008) noted that most
Dasypoda species seem to be oligolectic on actinomorphic
plant families (i.e. those with radiate symmetry). Dasypoda
hirtipes is the most widespread and abundant species
visiting different plant species in the families Apiaceae,
Asteraceae, Convolvulaceae, Dipsacaceae, Euphorbiaceae,
Fabaceae, Lamiaceae, and Scrophulariaceae. Field
observations and records reveal that it has a very strong
preference to C. intybus in nature; however, in various
localities, it also forages exclusively on Cephalaria gigantea,
C. procera, and Melilotus officinalis. Dasypoda pyriformis,
D. tubera, D. warnckei, D. toroki, and D. visnaga forage on
various plant species in different families, although they
exclusively visit the plant species in the family Asteraceae
for pollen. D. argentata, D. braccata, D. frieseana, D.
longigena, and D. spinigera forage mainly on the plant
species in the family Dipsacaceae, particularly the
Cephalaria species. I observed that in mountainous areas
of East Anatolia (Ağrı, Erzurum, and Kars provinces),
above 2000 m, almost all the plants of C. gigantea and C.
procera host 1 or more of the above-mentioned Dasypoda
species. Among the 2 Macropis species occurring in
Turkey, only 1 sample has been collected belonging to M.
frivaldszkyi, from Lysimachia vulgaris. Michez et al. (2008)
noted that Macropis bees are apparently all oligolectic on
Lysimachia. Females collect pollen and oil on Lysimachia,
whereas nectar is usually collected from a wide variety of
host plants.
Concerning the Melitta species visiting wild plants,
the present data revealed that although Melitta species
visit various plant species in different families, they show a
clear preference for the family Fabaceae. In particular, M.
leporina and M. dimidiata are widespread and abundant
bees, visiting various wild plants in the Fabaceae. Among
them, Astragalus, Lotus, and Melilotus species have special
importance due to occurring in mountainous sloping
areas, especially in eastern part of the country. These are
important pasture plants for grazing domestic animals as
well as wildlife.
Moreover, Melittidae species visit many plant species
in different families. Turkey is a mountainous and hilly
country. The average altitude is approximately 1250 m, and
more than 60% of the total land has more than 15% slopes.
Turkey is thus very sensitive to erosion and flood disasters.
It is well known that plants provide protective cover on the
land and prevent soil erosion. Diversity of various plants
species is associated with many kinds of pollinators. The
Melittidae include some of these pollinators.
I should admit that in Turkey some farmers and even
agricultural experts are unaware of the importance of
bees as pollinators of crops and wild plants. Most of them
rely on the honey bee alone (A. mellifera) for most crop
pollination while the majority believe that wild bees are
harmful insects, whereas, as we indicated previously,
pollinator bees are well known to provide key ecosystem
services to both natural and agroecosystems.
Acknowledgments
I would like to thank Dr Denis Michez (Université de Mons,
Laboratoire de Zoologie, Mons, Belgium) for determining
certain specimens. I also thank all my colleagues, Osman
Ecevit, Şaban Güçlü, Rüstem Hayat, Erol Yıldırım, İrfan
Aslan, Göksel Tozlu, Önder Çalmaşur, Hidayet Bostan,
Muhammed Atamanalp, Ümit Avcı, Engin Kılıç, Saliha
Çoruh, Memiş Kesdek, and Fatih Aydın, who helped me to
collect certain bee samples in the field, and İrfan Çoruh for
determining plant species. I thank İbrahim Yücel Özbek, who
kindly prepared the distribution maps. I am also thankful to
2 anonymous reviewers for their helpful comments.
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