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Hoplitis (Hoplitis) galichicae spec. nov., a new osmiine bee species from Macedonia with key to the European representatives of the Hoplitis adunca species group (Megachilidae, Osmiini)

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Hoplitis (Hoplitis) galichicae spec. nov., a new European osmiine bee species belonging to the Hoplitis adunca species group (Osmiini) is described and diagnosed. It is currently known only from the Galichica mountain range in southwestern Macedonia. Analysis of pollen contained in the metasomal scopae revealed that all females of the type series collected pollen on the flowers of Sedum (Crassulaceae), which is unexpected as most other members of the Hoplitis adunca species group are oligolectic or mesolectic on flowers of Boraginaceae and/or Fabaceae. An identification key including all European representatives of the Hoplitis adunca species group is given.
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Accepted by V. Gonzalez: 21 Mar. 2016; published: 13 May 2016
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2016 Magnolia Press
Zootaxa 4111 (2): 167
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http://www.mapress.com/j/zt/
Article
167
http://doi.org/10.11646/zootaxa.4111.2.5
http://zoobank.org/urn:lsid:zoobank.org:pub:CF3C0DBB-F5C5-41F3-B845-0CDD7E090F03
Hoplitis (Hoplitis) galichicae spec. nov., a new osmiine bee species from
Macedonia with key to the European representatives of the Hoplitis adunca
species group (Megachilidae, Osmiini)
ANDREAS MÜLLER
ETH Zurich, Institute of Agricultural Sciences, Biocommunication and Entomology, Schmelzbergstrasse 9/LFO, 8092 Zurich, Switzer-
land. E-mail: andreas.mueller@usys.ethz.ch
Abstract
Hoplitis (Hoplitis) galichicae spec. nov., a new European osmiine bee species belonging to the Hoplitis adunca species
group (Osmiini) is described and diagnosed. It is currently known only from the Galichica mountain range in southwestern
Macedonia. Analysis of pollen contained in the metasomal scopae revealed that all females of the type series collected
pollen on the flowers of Sedum (Crassulaceae), which is unexpected as most other members of the Hoplitis adunca species
group are oligolectic or mesolectic on flowers of Boraginaceae and/or Fabaceae. An identification key including all Eu-
ropean representatives of the Hoplitis adunca species group is given.
Key words: Apiformes, Galichica national park, host plant choice, Hymenoptera, Sedum
Introduction
Osmiine bees constitute a tribe within the family Megachilidae (Michener 2007; Praz et al. 2008). Including taxa
from the Canary Islands, Cyprus and the Caucasus, about 270 Osmiini species have been recorded so far in Europe
(Müller 2015). Recently, 15 new European osmiine bee species mainly from Spain, Sicily or Greece have been
described (Müller 2012). This indicates that the bee fauna of southwestern, southern and southeastern Europe is
still incompletely known and that remote and undersampled regions might harbour hitherto overlooked species,
even of the Megachilidae, which are among the best studied bee taxa in Europe. Recent examination of megachilid
bee material from Macedonia revealed the existence of a further undescribed osmiine bee species, which belongs to
the Hoplitis adunca species group of Hoplitis (Hoplitis). In the present publication, this new species is described
and diagnosed against its closest relatives. In addition, a key is given for all 25 European representatives of the
Hoplitis adunca species group, which is one of the taxonomically most challenging osmiine bee taxon due to the
high morphological uniformity among its species, especially in the female sex.
Material and methods
Morphological terminology and definitions for body measurements follow Michener (2007) with the following
specifications, which refer to the species description and the identification key: i) the distance between lateral
ocellus and preoccipital ridge was measured in top view rather than in lateral view; ii) the diameter of an ocellus
was measured under inclusion of the ocellar border, which is often of the same colour as the surrounding cuticle
thereby differing from the usually light colour of the central part of the ocellus; iii) the length of a segment of the
labial palpus was measured from its sclerotized base to the sclerotized base of the subsequent segment; iv) the
length of an antennal segment was measured along its lower margin, while its width corresponds to the maximal
width of the segment. Numbering of antennal segments starts from the scape, which is antennal segment 1. The
number of a segment belonging to a segmented body part is put into parentheses if a character of that segment is
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not developed in all individuals; thus, “terga 1–4(5) with apical white hair bands” means that tergum 5 sometimes
lacks apical hair bands. Measurements to the nearest 0.1mm or 0.5mm (for body length) were taken using an ocular
micrometer on an Olympus VMT stereomicroscope. Photomicrographs were taken with the digital microscope
Keyence VHX-2000. To assess the pollen hosts of the species, scopal pollen contents of all available females were
analysed by light microscopy applying the method of Sedivy et al. (2013b).
Species description
Hoplitis (Hoplitis) galichicae Müller spec. nov.
Holotype. MACEDONIA: N.P. Galicica, 600–1500m, 26.6.2014, ♂ (leg. J. Halada & M. Fabianová). Deposited in
the private collection of M. Schwarz (Ansfelden).
Paratypes. MACEDONIA: N.P. Galicica, 600–1500m, 26.6.2014, 25♀, 6♂ (leg. J. Halada & M. Fabianová).
Deposited in the Entomological Collection of ETH Zurich and the private collections of M. Schwarz (Ansfelden)
and the author.
Diagnosis. With a body length of only 5–6.5mm, H. galichicae is the smallest representative of the Hoplitis
adunca species group in Europe, where all other species of that group attain a body length of at least 6–6.5mm. The
female is further characterized by the only weakly convex clypeus (Fig. 1) in combination with the long and
slender, almost straight and apically pointed hind tibial spurs (Fig. 3), the short antennal segment 3, which is about
1.5x as long as wide, and the sparse pilosity of the supraclypeal area (Fig. 1). The male is additionally characterized
by the oval, laterally rounded and evenly haired lobes of the membraneous appendage of sternum 6, which are
directed laterally and separated from each other by an angle of almost 120
o
(Fig. 6), in combination with the
slender, almost straight and apically pointed hind tibial spurs.
Description. FEMALE: Body length 5–6.5mm. Head: Head about 0.95x as long as wide (Fig. 1). Distance
between lateral ocellus and preoccipital ridge about 1.6x as long as ocellar diameter. Maximal width of genal area
about 0.6x as long as maximal width of compound eye. Second segment of labial palpus 1.75–1.85x as long as first
segment. Antennal segment 3 about 1.5x as long as wide. Clypeus weakly convex (Fig. 1); in profile, clypeal
surface not or only slightly projecting above surface of supraclypeal area. Punctation of clypeus and supraclypeal
area very dense with interspaces rarely exceeding the diameter of half a puncture except sometimes for a small
polished area at the clypeal base (Fig. 1). Face rather sparsely covered with long white hairs, which do not hide the
cuticular surface neither on clypeus and paraocular nor on supraclypeal area (Fig. 1). Mesosoma: Punctation of
scutum and scutellum dense with interspaces usually not exceeding the diameter of one puncture except for the
lateral parts of the scutum, where the interspaces may reach the diameter of one and a half to two punctures.
Punctation of mesepisternum dense with interspaces varying in size between the diameter of half a puncture and
the diameter of one to one and a half punctures. Tibial spur of fore leg with short and pointed tooth, which is about
as long as its basal width. Inner tibial spur of hind leg yellowish and long and slender, its apex pointed and only
very slightly curved (Fig. 3). Pilosity on inner side of hind basitarsus yellowish-white. Tegula blackish in its
anterior third and yellowish-brown in its posterior two thirds. Stigma and veins of fore wing (dark) brown to black.
Metasoma: Punctation of tergal discs rather scattered with interspaces varying in size between the diameter of one
and a half to one puncture (laterally) and the diameter of two to three, rarely four punctures (medially). Punctation
of marginal zones of terga distinctly finer and denser than on discs and restricted to the basal half. Terga 1–4(5)
with short apical white hair bands, which are medially interrupted in older specimens. Longest hairs on median half
of tergum 1 less than half as long as maximal length of lateral hair tuft. Discs of terga 5–6 with very sparse
appressed white pilosity. Sternum 6 densely punctured and dull except usually for a narrow unpunctured median
zone; its apical carina very weak, evenly rounded, of constant height throughout and continuous or only narrowly
interrupted medially (Fig. 2). Scopa whitish except sometimes for some blackish hairs at its base.
MALE: Body length 5–6mm. Head: Head about 0.85x as long as wide. Distance between lateral ocellus and
preoccipital ridge about 1.45x as long as ocellar diameter. Maximal width of genal area about 0.5x as long as
maximal width of compound eye. Second segment of labial palpus 1.7–1.8x as long as first segment. Antennal
segment 3 1.1x, segment 4 0.7x, segments 5–11 0.75x, segment 12 1x and segment 13 1.6x as long as wide (Fig. 4).
Anterior side of antennal segments (3)4–12 and posterior side of segments (5)6–12 yellowish except sometimes for
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NEW HOPLITIS FROM MACEDONIA
the slightly darkened upper margin (Fig. 4). Mesosoma: Punctation of scutum, scutellum and mesepisternum
similar to that of the female albeit slightly finer and less dense on scutum and scutellum. Tibial spurs of hind leg
yellowish and slender, almost straight and apically pointed. Colour of tegula and of wing venation as in the female.
Metasoma: Punctation of terga similar to that of the female. Terga 1–4(5) with short apical white hair bands, which
are medially interrupted in older specimens. Apical margin of tergum 6 medially crenulate and laterally with
distinct tooth (Fig. 5). Apical margin of tergum 7 evenly rounded (Fig. 5). Sterna 2–3 with strong transverse
swellings, which are almost devoid of punctures and narrowly interrupted medially (Fig. 7). Transverse swelling of
sternum 4 distinctly less strongly developed than on sterna 2–3, sparsely punctured and broadly interrupted
medially (Fig. 7). Apical margins of sterna 2–4 almost straight and beset with white hairs, which are much longer
laterally than medially (Fig. 7). Sternum 5 strongly shagreened in its basal half, rather coarsely and densely
punctured in its preapical part with interspaces reaching the diameter of up to one and a half to two punctures, and
very finely and densely punctured along its marginal zone; its apical margin slightly emarginated medially (Fig. 6).
Base of sternum 6 with a pair of large translucent flaps (Fig. 6). Lateral lobes of the bilobed membraneous
appendage of sternum 6 oval (about 1.6x as wide as long), laterally rounded to truncate, densely covered with hairs
slightly directed backwards and separated from each other by an angle of almost 120
o
(Fig. 6).
Distribution. Known so far only from the Galichica national park in southwestern Macedonia.
FIGURE 1–7. Hoplitis galichicae. 1: Head of female. 2: Sterna 5–6 of female. 3: Inner hind tibial spur of female. 4: Antenna
of male. 5: Terga 4–7 of male. 6: Sterna 5–6 of male. 7: Sterna 2–5 of male.
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Pollen hosts. The pollen loads of six females of the type series all consisted exclusively of pollen of an
unknown species of Sedum (Crassulaceae). Interestingly, the pollen masses in the metasomal scopae of all females
were noticeably moist, indicating that the females had added nectar to the collected pollen probably in order to
facilitate the transport of the minute Sedum pollen grains, which were found to have a diameter of only about
15μm. If H. galichicae should turn out to have a clear or even exclusive preference for the flowers of Crassulaceae
as pollen hosts, it would be an exception within the Hoplitis adunca species group as most of its members are
oligolectic or mesolectic on flowers of Boraginaceae (mostly Echium) and/or Fabaceae (Sedivy et al. 2013b;
Müller 2015; Tab. 1). However, Crassulaceae pollen was found to be rarely collected also by H. loti and H. ravouxi,
two mesolectic species of the Hoplitis adunca species group, which otherwise show a clear preference for the
pollen of Loteae (Fabaceae) (Sedivy et al. 2013b).
Nesting biology. Unknown. As most species of the Hoplitis adunca species group nest in depressions or
cavities of stones and rocks and build their brood cells with mud often combined with small pebbles (Sedivy et al.
2013a; Müller 2015), H. galichicae is expected to exhibit an analogous nesting behaviour as its relatives.
Etymology. galichicae = from the Galichica mountain range in southwestern Macedonia.
Key to the European species of the Hoplitis adunca species group
Based on a molecular phylogeny of the genus Hoplitis, Sedivy et al. (2013b) merged the former Hoplitis subgenera
Annosmia Warncke, Bytinskia Mavromoustakis, Coloplitis Griswold and Hoplitis Klug into a single subgenus
Hoplitis. This enlarged taxon is currently divided into five species groups (Müller 2015). Two of these groups
occur in Europe, i.e. the Hoplitis adunca species group and the Hoplitis annulata species group, which correspond
to the former subgenera Hoplitis and Annosmia, respectively (Michener 2007). The males of these two groups
distinctly differ by i) the shape of tergum 7, which is evenly rounded to truncate in the Hoplitis adunca species
group but deeply bifid in the Hoplitis annulata species group, ii) the form of the mandible, which is usually two-
toothed in the former but always three-toothed in the latter, and iii) the shape of the last antennal segment, which is
apically hooked down in the Hoplitis annulata species group but only rarely so in the Hoplitis adunca species
group. In contrast, no clear morphological characters are known that unambiguously separate the females of the
two species groups. The following identification key comprises all 25 species of the Hoplitis adunca species group
known to occur in Europe including the Canary Islands and Cyprus but excluding the Caucasus.
Compared to the males, which are rather well characterized by the shape of both the antenna and the
membraneous appendage of sternum 6, the females of the Hoplitis adunca species group are morphologically
highly uniform and notoriously difficult to identify. To facilitate identification, the European distribution area and
the pollen host preferences are detailed for each species in Table 1. However, our present knowledge on the
distribution of numerous species is far from complete and might partly even contain errors. In fact, due to the
difficult identification of numerous species, which has resulted in many misidentifications on the one hand and in
substantial amounts of undetermined material in collections on the other hand, much work is still needed to clarify
the exact distribution of the representatives of the Hoplitis adunca species group in Europe. The key is tailored to
European populations. As North African and Asian populations of several species slightly differ morphologically
from those of Europe, the proper identification of females from outside Europe might not always be possible.
Two females and two males of Hoplitis adunca all collected at the same day and locality in western Sardinia
are unusually small and have a shorter proboscis and a narrower vertex compared to mainland individuals of H.
adunca, from which they otherwise do not differ morphologically. Similarly, a single Hoplitis male from Crete
differs from males of Hoplitis pici, which it otherwise closely resembles, by the shape of both the genitalia and the
last antennal segment. More material from Sardinia and Crete is needed to judge whether these specimens are
merely aberrant specimens of H. adunca and H. pici, respectively, or whether they should be assigned to new
(sub)species. These two still enigmatic taxa are not included in the key.
Females
1 Galea of proboscis with specialized pollen-harvesting bristles, which are curved at an angle of 45–90
o
in their apical third. Tib-
ial spur of fore leg apically truncate or rounded, without projecting tooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- Galea of proboscis with unspecialized bristles, which are straight along their whole length. Tibial spur of fore leg apically pro-
longed into a shorter or longer tooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2(1) Curved bristles limited to the basal half of the galea. Tibial spur of fore leg apically truncate. Body length 8–9mm . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis lithodorae Müller
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NEW HOPLITIS FROM MACEDONIA
- Curved bristles covering the whole galea except for a short apical zone. Tibial spur of fore leg apically rounded. Body length
7.5–9mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis pici (Friese)
3(1) Clypeus densely covered with specialized suberect pollen-harvesting bristles, which have a strong basal part of brownish
colour and a thin and slightly wavy apical part of whitish colour. Punctation of clypeus coarse and rather scattered with inter-
spaces reaching the diameter of one puncture. Body length 6–7mm . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis bihamata (Costa)
- Clypeus covered with unspecialized whitish pilosity. Punctation of clypeus fine and dense with interspaces usually not reach-
ing the diameter of one puncture except for the median or basal part of the clypeus in some species, where the punctation is
more scattered . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4(3) Lateral lobes of pronotum distinctly inflated and sparsely punctured with polished interspaces. Punctation of anterior half of
scutum on both sides of the median impression scattered with interspaces reaching the diameter of up to one and a half to two
punctures. Punctation between lateral ocellus and compound eye scattered with interspaces reaching the diameter of up to two,
rarely more punctures. Pilosity of mesosoma and terga very long and scrubby. Body length 8.5–10.5mm . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis strymonia Tkalcu
- Lateral lobes of pronotum not inflated and usually densely punctured and dull. Punctation of anterior half of scutum on both
sides of the median impression as well as between lateral ocellus and compound eye usually dense with interspaces rarely
exceeding the diameter of one puncture except for the Canarian endemite H. perambigua, where the punctation is less dense.
Pilosity of mesosoma and terga often shorter and less scrubby . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5(4) Proboscis very long, second segment of labial palpus about 2.9x as long as first segment; in repose, proboscis slightly sur-
passes trochanter of hind leg. Body length 6.5–8mm . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis holmboei (Mavromoustakis)
- Proboscis shorter, second segment of labial palpus less than 2.5x as long as first segment; in repose, proboscis does not reach
trochanter of hind leg . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6(5) Apical margin of sternum 6 medially prolonged into a distinct and well delimited tooth of narrowly triangular to linear shape.
Body length often exceeding 8–9mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
- Apical margin of sternum 6 medially rounded to tapering, but never with distinct and well delimited tooth. Body length usually
less than 9–10mm except for H. lepeletieri . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .13
7(6) Sternum 6 lateroapically without or with very weak carina, almost plane and densely punctured and dull. Antennal segment 3
more than 2.5x as long as wide. Discs of terga 2–5 with long pilosity. Body length 10.5–12.5mm . . Hoplitis mucida (Dours)
- Sternum 6 lateroapically with very strong carina, zone adjacent to the carina sparsely punctured and polished. Antennal seg-
ment 3 about 2x as long as wide. Discs of terga 2–5 with short pilosity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8(7) Pilosity of inner side of basitarsus of hind leg brownish at least apically. Scopa often more or less smoky brown. When seen
from behind, longest hairs on median half of tergum 1 almost as long as maximal length of lateral hair tuft. Inner tibial spur of
hind leg apically distinctly curved (only slightly curved in north African specimens). Apical tooth of tibial spur of fore leg
about as long as its basal width (often longer in north African specimens) and acute. Body length 8–10.5mm . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis fertoni (Pérez)
- Pilosity of inner side of basitarsus of hind leg yellowish-white to yellowish-red. Scopa whitish except for its base, which is
occasionally smoky brown. When seen from behind, longest hairs on median half of tergum 1 distinctly shorter than maximal
length of lateral hair tuft. Inner tibial spur of hind leg apically only slightly curved. Apical tooth of tibial spur of fore leg longer
than its basal width or rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9(8) Apical tooth of tibial spur of fore leg more or less rounded. Punctation of lateral parts of clypeus less dense with distinct pol-
ished interspaces. Punctation of supraclypeal area scattered with interspaces reaching the diameter of one to one and a half,
rarely two punctures. Second segment of labial palpus 1.5–1.6x as long as first segment. Body length 8–10mm. No morpholog-
ical characters are known to distinguish the following two species. However, identification should be possible based on the
apparently almost non-overlapping distribution areas (see Tab. 1) . . Hoplitis fabrei Zanden and Hoplitis pallicornis (Friese)
- Apical tooth of tibial spur of fore leg acute. Punctation of lateral parts of clypeus very dense lacking distinct polished inter-
spaces. Punctation of supraclypeal area dense with interspaces rarely reaching the diameter of one puncture. Second segm ent
of labial palpus at least 1.7x as long as first segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10(9) Distance between lateral ocellus and preoccipital ridge more than 3x as long as ocellar diameter. Tibial spurs of hind leg red-
dish-brown to black. Body length 10.5–13.5mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis manicata (Morice)
- Distance between lateral ocellus and preoccipital ridge less than 3x as long as ocellar diameter. Tibial spurs of hind leg black,
reddish-brown or yellowish-red. Body length rarely exceeding 10.5mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11(10) Distance between lateral ocellus and preoccipital ridge 1.8–2x as long as ocellar diameter. Second segment of labial palpus
1.7–1.8x as long as first segment. Punctation of discs of terga 1–3(4) rather dense: largest interspaces on the two longitudinal
fifths adjacent to each side of the median fifth rarely exceeding the diameter of one and a half punctures. Tibial spurs of hind
leg reddish-brown to black. Body length 8–10.5mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis benoisti (Alfken)
- Distance between lateral ocellus and preoccipital ridge at least 2.4x as long as ocellar diameter. Second segment of labial pal-
pus at least 2.2x as long as first segment. Punctation of discs of terga 1–3(4) very scattered: largest interspaces on the two lon-
gitudinal fifths adjacent to each side of the median fifth usually reaching the diameter of three to four, sometimes even more
punctures. Tibial spurs of hind leg black or yellowish-red . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12(11) Tibial spurs of hind leg mostly black to dark reddish-brown. Outer marginal zone of tegula without punctures on its posterior
half or with broadly interrupted row(s) of punctures. Second segment of labial palpus slightly less than 2.3x as long as first
segment. Interspaces between punctures on vertex usually without or with few micropunctures. Marginal zones of terga mostly
black. Body length 9–11mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis adunca (Panzer)
- Tibial spurs of hind leg yellowish-red. Outer marginal zone of tegula often punctured along its whole length. Second segment
of labial palpus slightly more than 2.3x as long as first segment. Interspaces between punctures on vertex usually with many
micropunctures. Marginal zones of terga often more or less reddish-brown. Body length 9–11mm . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Hoplitis submanicata Zanden
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13(6) Apex of inner tibial spur of hind leg acute and almost straight to slightly bent at an angle of less than 45
o
. . . . . . . . . . . . . . 14
- Apex of inner tibial spur of hind leg blunt or strongly curved at an angle of more than 45
o
. . . . . . . . . . . . . . . . . . . . . . . . . . 19
14(13) Antennal segment 3 longer, 1.75–2x as long as wide. When seen from behind, longest hairs on median half of tergum 1 almost
as long as maximal length of lateral hair tuft . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
- Antennal segm ent 3 shorter, about 1.5x as long as wide. When seen from behind, longest hairs on median half of tergum 1
about half as long as maximal length of lateral hair tuft or shorter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
15(14) Pilosity of inner side of hind basitarsus (dark) greyish-brown. Apical white hair bands of terga 1–4 broadly interrupted. Punc-
tation of anterior half of scutum moderately dense with interspaces often reaching or exceeding the diameter of one puncture.
Punctation of median half of tergal discs 1–3 very fine and widely scattered. Body length 6–7.5mm . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis perambigua (Peters)
- Pilosity of inner side of hind basitarsus yellowish-white. Apical white hair bands of terga 1–4 not interrupted in fresh speci-
mens. Punctation of anterior half of scutum very dense with interspaces rarely exceeding the diameter of half a puncture. Punc-
tation of median half of tergal discs 1–3 distinctly coarser and denser. Body length at least 8mm . . . . . . . . . . . . . . . . . . . . . 16
16(15) Pilosity of terga shorter: when seen in profile, longest hairs on median half of tergal disc 2 about 2x as long as maximal width
of antennal flagellum, on tergal discs 3–4 less than 1.5x as long as maximal width of antennal flagellum. Sternum 6 lateroapi-
cally usually with weak carina. Body length 8–9mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis anthocopoides (Schenck)
- Pilosity of terga longer: when seen in profile, longest hairs on median half of tergal disc 2 about 3x as long as maximal width
of antennal flagellum, on tergal discs 3–4 about 2x as long as maximal width of antennal flagellum. Sternum 6 lateroapically
usually with rather strong carina. Body length 9.5–11mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis lepeletieri (Pérez)
17(14) Clypeus weakly convex (Fig. 1); in profile, clypeal surface not or only slightly projecting above surface of supraclypeal area.
Supraclypeal area with sparse pilosity, which never hides its surface (Fig. 1). Body length 5–6.5mm . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis galichicae spec. nov.
- Clypeus strongly convex; in profile, clypeal surface distincly projecting above surface of supraclypeal area. Supraclypeal area
with rather dense pilosity, which partly hides its surface. Body length at least 6.5mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
18(17) Apical tooth of tibial spur of fore leg as long as its basal width or shorter. Distance between lateral ocellus and preoccipital
ridge about 1.5x as long as ocellar diameter. Second segment of labial palpus about 1.4x as long as first segment. Head slightly
shorter than wide. Carina along apical margin of sternum 6 weakly developed, medially usually not interrupted and here only
slightly lower than laterally. Punctation of mesepisternum and mesosternum moderately dense with interspaces often exceed-
ing the diameter of one puncture. Body length 6.5–7.5mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis hilbera Müller
- Apical tooth of tibial spur of fore leg longer than its basal width. Distance between lateral ocellus and preoccipital ridge
1.8–1.9x as long as ocellar diameter. Second segment of labial palpus about 1.7x as long as first segment. Head slightly longer
than wide. Carina along apical margin of sternum 6 well developed, medially usually interrupted or laterally distinctly higher
than medially. Punctation of mesepisternum and mesosternum dense with interspaces rarely exceeding the diameter of one
puncture. Body length 7–8mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis marchali (Pérez)
19(13) Inner tibial spur of hind leg short and stout, 4–5x as long as its maximal width (including marginal teeth). Teeth along upper
margin of inner hind tibial spur distinct, rather long and well separated from each other. Hind tibial spurs yellowish. Clypeus
medially with very sharp and narrow longitudinal carina. Second segment of labial palpus about 1.4x as long as first segment.
Anterior side of antennal segments (5,6)7–11(12) light reddish-brown. Body length 6.5–8mm . . . Hoplitis carinata (Stanek)
- Inner tibial spur of hind leg longer, at least 6x as long as its maximal width (including marginal teeth). Teeth along upper mar-
gin of inner hind tibial spur often indistinct, short and not well separated from each other. Hind tibial spurs yellowish or black-
ish. Clypeus medially either with or without longitudinal carina or polished midline. Second segment of labial palpus at least
1.5x as long as first segment. Anterior side of antennal segments usually blackish, some segments occasionally partly dark red-
dish-brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
20(19) Punctation of anterior third of scutum fine and very dense with interspaces usually smaller than the diameter of half a puncture.
Disc of tergum 5 covered with rather dense and appressed whitish pilosity. Pilosity of terga longer: when seen in profile, lon-
gest hairs on median half of tergal discs 2–4 1.3–1.5x as long as maximal width of antennal flagellum. Body length 7–9mm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis loti (Morawitz)
- Punctation of anterior third of scutum coarser and less dense with interspaces usually reaching the diameter of half a puncture
to one puncture. Disc of tergum 5 without dense and appressed whitish pilosity. Pilosity of terga shorter: when seen in profile,
longest hairs on median half of tergal discs 2–4 at most as long as maximal width of antennal flagellum . . . . . . . . . . . . . . 21
21(20) Hind tibial spurs dark brown to black. Apex of inner hind tibial spur almost blunt and bent at right angles. Second segment of
labial palpus 1.7–1.8x as long as first segment. Body length 6.5–7.5mm . . . . . . . . . . . . . Hoplitis insularis (Schmiedeknecht)
- Hind tibial spurs yellowish. Apex of inner hind tibial spur either almost blunt and bent at right angles or tapering and evenly
curved. Second segment of labial palpus at most 1.7x as long as first segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
22(21) Clypeus strongly convex; in profile, clypeal surface distincly projecting above surface of supraclypeal area. Apex of inner hind
tibial spur regularly tapering, evenly curved at an angle of 50–60
o
and rather long. Second segment of labial palpus about 1.7x
as long as first segment. Clypeus (always?) with rather indistinct and more or less polished longitudinal midline. Body length
7–7.5mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis ochraceicornis (Ferton)
- Clypeus weakly convex; in profile, clypeal surface only slightly projecting above surface of supraclypeal area. Apex of inner
hind tibial spur more or less blunt, abruptly curved at an angle of 60–90
o
and usually very short. Second segment of labial pal-
pus 1.5–1.6x as long as first segment. Clypeus medially either with or without polished longitudinal midline or carina . . . 23
23(22) Clypeus usually densely punctured throughout, sometimes with indistinct polished longitudinal midline on its basal half. Body
length 6.5–7.5mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis jheringii (Ducke)
- Clypeus usually with distinct, uninterrupted and polished longitudinal midline or carina. Body length 7–8.5mm . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis ravouxi (Pérez)
Zootaxa 4111 (2) © 2016 Magnolia Press
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173
NEW HOPLITIS FROM MACEDONIA
Males
1 Sternum 6 without membraneous appendage clearly separated from its apical margin, but with almost hairless medioapical
impression of roughly triangular shape . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- Sternum 6 with membraneous appendage at its apical margin, which is usually bilobed and densely haired . . . . . . . . . . . . . 5
2(2) Medioapical impression of sternum 6 neither with preapical hair pencil nor median projection, its punctation mediobasally
coarse and extremely dense without any interspaces. Body length 8–10mm . . . . . . . . . . . . . . . . . . Hoplitis benoisti (Alfken)
- Medioapical impression of sternum 6 with preapical hair pencil or median projection, its punctation lacking or very fine . . 3
3(2) Medioapical impression of sternum 6 with long preapical hair pencil projecting at right angles from the sternal surface. Apical
margin of medioapical impression of sternum 6 medially prolonged into a long and narrow process. Body length 9–12mm . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis adunca (Panzer)
- Medioapical impression of sternum 6 with narrow median projection, which is formed by two longitudinal and apically fused
keels and consists of a lower anterior and a higher posterior part. Apical margin of medioapical impression of sternum 6 medi-
ally emarginate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4(3) Posterior part of median projection of sternum 6 about three times as high as anterior part, its maximal height roughly equal-
ling length of anterior part. Distance between lateral ocellus and preoccipital ridge at least 3x as long as ocellar diameter.
Preapical lateral swellings of sternum 5 weakly developed and rounded. Body length 12–15mm . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Hoplitis manicata (Morice)
- Posterior part of median projection of sternum 6 about twice as high as anterior part, its maximal height distinctly shorter than
length of anterior part. Distance between lateral ocellus and preoccipital ridge at most 2.5x as long as ocellar diameter. Preapi-
cal lateral swellings of sternum 5 well developed and edged. Body length 10–12mm . . . . . . . . Hoplitis submanicata Zanden
5(1) Apex of last antennal segment curved downwards . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
- Apex of last antennal segment not curved downwards . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
6(5) Last antennal segment apically with short projection of roughly triangular shape, which is directed downwards. Apical margin
of tergum 6 with two triangular teeth on each side. Antennal segment 3 distinctly longer than segment 4. Body length
10.5–12.5mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis mucida (Dours)
- Last antennal segment narrowed towards its apex, which is pointed and curved downwards. Apical margin of tergum 6 with
one triangular tooth on each side. Antennal segment 3 slightly shorter than segment 4. Body length 7–9mm . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis pici (Friese)
7(5) Lower margin of antennal segment 3 apically broadened and detached from base of segment 4. Membraneous appendage at
apical margin of sternum 6 medially without uninterrupted transverse hair stripe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
- Lower margin of antennal segment 3 neither broadened nor detached from base of segment 4; if slightly detached, membrane-
ous appendage at apical margin of sternum 6 medially with uninterrupted transverse hair stripe . . . . . . . . . . . . . . . . . . . . . . 9
8(7) Antennal segment 4 as long as and larger than segment 3, about three quarters as long as wide. Membraneous appendage at
apical margin of sternum 6 consisting of two roughly triangular lobes densely covered with appressed long pilosity, which is
directed inwards and completely conceals the membraneous surface. Inner spur of hind tibia stout, its apex short and strongly
curved. Body length 7–8.5mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis carinata (Stanek)
- Antennal segment 4 distinctly shorter and smaller than segment 3, about half as long as wide. Membraneous appendage at api-
cal margin of sternum 6 consisting of two roughly quadrangular lobes beset with inconspicuous short hairs, which are suberect
and do not completely conceal the membraneous surface. Inner spur of hind tibia slender, its apex long and slightly curved.
Body length 6.5–7.5mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis ochraceicornis (Ferton)
9(7) Membraneous appendage at apical margin of sternum 6 V-shaped, i.e. consisting of two slender lobes, which are at least twice
as long as wide and only basally fused; these lobes are directed backwards rather than laterally and diverge from each other by
an angle of less than 90
o
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
- Membraneous appendage at apical margin of sternum 6 of other shape . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
10(9) Apical margin of sternum 6 medially with small roundish excision. Membraneous appendage of sternum 6 mediobasally with-
out roundish tuft of hairs. Lower half of anterior side of antennal segments (3)4–12(13) and posterior side of antennal segments
(6)7–12(13) yellowish. Last antennal segment slightly twisted with upper and lower margin being in different planes. Body
length 8–10mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis fertoni (Pérez)
- Apical margin of sternum 6 deeply and widely excised. Membraneous appendage of sternum 6 mediobasally with roundish tuft
of hairs. Antenna uniformly dark except sometimes for its lower margin, which is partly yellowish-brown. Last antennal seg-
ment not modified . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11(10) Apical margin of sternum 5 laterally emarginate. Preapical swellings of sterna 2–4 strongly developed, transition between their
anterior and posterior part edged. Body length 10–11.5mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis lepeletieri (Pérez)
- Apical margin of sternum 5 laterally evenly rounded. Preapical swellings of sterna 2–4 less strongly developed, transition
between their anterior and posterior part rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12(11) Pilosity of inner side of hind basitarsus yellowish-white. Antennal segment 3 about 1.7x as long as segment 4. Punctation of
scutum dense with interspaces usually smaller than the diameter of one puncture on each side of the medioapical suture. Punc-
tation of terga rather coarse and dense with interspaces rarely exceeding the diameter of two to three punctures on tergal sides.
Body length 7.5–10mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis anthocopoides (Schenck)
- Pilosity of inner side of hind basitarsus (dark) greyish-brown. Antennal segment 3 about 1.4x as as long as segment 4. Puncta-
tion of scutum less dense with interspaces usually exceeding the diameter of one puncture on each side of the medioapical
suture. Punctation of terga very fine and scattered with interspaces reaching the diameter of four to five punctures on tergal
sides. Body length 6–7.5mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis perambigua (Peters)
13(9) Antennal segment 3 distinctly longer than wide. Membraneous appendage of sternum 6 consisting of a roughly quadrangular
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Zootaxa 4111 (2) © 2016 Magnolia Press
plate, which bears apicolaterally two oval lobes beset with long hairs. Apical margin of tergum 7 medially with small but dis-
tinct triangular emargination. Pilosity of mesosoma and terga very long and tousled. Terga without dense, short and apressed
apical white hair bands. Body length 7.5–8mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis strymonia Tkalcu
- Antennal segment 3 as long as wide or shorter; if slightly longer than wide, membraneous appendage of sternum 6 of other
shape. Apical margin of tergum 7 evenly rounded or truncate, rarely with very shallow and inconspicuous median emargin-
ation. Pilosity of mesosoma and terga shorter and less tousled. Terga with dense, short and appressed albeit often interrupted
apical white hair bands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14(13) Membraneous appendage of sternum 6 medially with uninterrupted and well limited transverse stripe of brownish hairs and
medioapically with small and densely haired buttonlike projection. Second last antennal segment with slightly converging
margins, last antennal segment tapering towards the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
- Membraneous appendage of sternum 6 neither with uninterrupted transverse stripe of brownish hairs nor medioapically with
densely haired buttonlike projection. Last two antennal segments usually almost parallel-sided . . . . . . . . . . . . . . . . . . . . . . 16
15(14) Antennal segment 4 about 0.8x as long as wide, slightly shorter than segment 3 and as long as segment 5. Body length
8–10mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis fabrei Zanden
- Antennal segment 4 about 0.6x long as wide, shorter than segments 3 and 5. Body length 8–10mm . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis pallicornis (Friese)
16(14) Lateral lobes of the bilobed membraneous appendage of sternum 6 of oval to oblong shape (i.e. distinctly wider than long ) and
separated from each other by an angle of 90
o
or more . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
- Lateral lobes of the bilobed membraneous appendage of sternum 6 of roundish to quadrangular shape (i.e. about as wide as
long or slightly longer) and separated from each other by an acute angle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
17(16) Spurs of hind tibia short and stout, almost blunt and usually of dark brown to blackish colour. Lateral margin of sternum 6 with
conspicuous bristles, which are directed backwards and about as long as maximal width of lateral lobe of membraneous
appendage of sternum 6. Body length 7–8.5mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis insularis (Schmiedeknecht)
- Spurs of hind tibia long and slender, pointed and of yellowish-brown to yellowish-white colour. Lateral margin of sternum 6
with inconspicuous bristles, which are not distinctly directed backwards and less than half as long as maximal width of lateral
lobe of membraneous appendage of sternum 6 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
18(17) Proboscis very long, second segment of labial palpus about 2.8x as long as first segment; in repose, proboscis reaches tro-
chanter of hind leg. Ventral surface of mesosoma between fore and mid coxa sparsely haired. Body length 6.5–8mm . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis holmboei (Mavromoustakis)
- Proboscis shorter, second segment of labial palpus 1.6–1.8x as long as first segment; in repose, proboscis does not reach tro-
chanter of hind leg. Ventral surface of mesosoma between fore and mid coxa with dense white pilosity . . . . . . . . . . . . . . . 19
19(18) Lateral lobes of membraneous appendage of sternum 6 laterally rounded to truncate and of regularly oval shape (Fig. 6). Inner
spur of hind tibia almost straight. Body length 5–6mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis galichicae spec. nov.
- Lateral lobes of membraneous appendage of sternum 6 laterally prolonged into a distinct and more or less acute tip. Inner spur
of hind tibia usually slightly to distinctly curved apically . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
20(19) Antennal segments 4–5 0.8–0.9x as long as wide with straight lower margin. Posterior side of antennal segments 8–10 with
distinct tubercle. Sternum 5 more or less densely punctured in front of preapical swelling. Lateral margin of sternum 6 apically
rounded. Body length 7.5–10mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis loti (Morawitz)
- Antennal segments 4–5 about 0.6x as long as wide with convex lower margin. Posterior side of antennal segments 8–10 almost
flat. Sternum 5 almost unpunctured in front of preapical swelling. Lateral margin of sternum 6 apically toothed. Body length
6.5–9mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis ravouxi (Pérez)
21(16) Antennal segment 4 shorter than segment 3. Pilosity along hypostomal carina uniformly short and sheared in appearance. Body
length 6.5–7mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis bihamata (Costa)
- Antennal segment 4 as long as antennal segment 3 or longer. Pilosity along hypostomal carina more irregular and increasing in
length towards foramen magnum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
22(21) Antennal segment 4 about 0.75x as long as wide. Preapical swellings of sternum 5 well developed and almost unpunctured.
Impressed area behind preapical swellings of sternum 5 medially sparsely punctured with interspaces reaching the diameter of
up to three punctures. Body length 7–9mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis marchali (Pérez)
- Antennal segment 4 as long as wide or slightly longer. Preapical swellings of sternum 5 weakly developed and moderately
dense to densely punctured. Impressed area behind preapical swellings of sternum 5 medially densely punctured with inter-
spaces not exceeding the diameter of one to two punctures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
23(22) Second segment of labial palpus about 1.75x as long as first segment. Anterior side of antennal segments 4–12 yellowish in its
lower third and blackish in its upper two thirds, the two colours being sharply separated. Posterior side of antennal segments
8–12 entirely yellowish. Body length 7–9mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis lithodorae Müller
- Second segment of labial palpus at most 1.6x as long as first segment. Antenna dark to reddish-brown, neither with sharp sep-
aration between yellowish and black colour on its anterior side nor with entirely yellowish coloured segments on its posterior
side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
24(23) Lateral lobes of membraneous appendage of sternum 6 laterally rounded. Apex of inner spur of hind tibia distinctly curved.
Apical margin of sternum 6 laterally with narrow pencil of hairs. Punctation of mesopleura dense with interspaces rarely
exceeding the diameter of one puncture. Body length 6–7mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplitis jheringii (Ducke)
- Lateral lobes of membraneous appendage of sternum 6 laterally extended into a short rounded tip. Apex of inner spur of hind
tibia very slightly curved. Apical margin of sternum 6 laterally without narrow pencil of hairs. Punctation of mesopleura less
dense with interspaces often exceeding the diameter of one puncture. Body length 6–8mm . . . . . . . . Hoplitis hilbera Müller
Zootaxa 4111 (2) © 2016 Magnolia Press
·
175
NEW HOPLITIS FROM MACEDONIA
TABLE 1. European distribution area and pollen host preferences of the 25 European representatives of the Hoplitis
adunca species group. Information on distribution and pollen hosts are based on Müller (2006, 2012, 2015, and
references therein) and Sedivy et al. (2013b). Definitions for categories of pollen host range are according to Cane &
Sipes (2006) and Müller & Kuhlmann (2008).
Species European distribution area Pollen host preferences
Hoplitis adunca (Panzer) entire Europe except for Great Britain, Ireland and
Scandinavia
oligolectic on Echium (Boraginaceae)
Hoplitis anthocopoides
(Schenck)
entire Europe except for Great Britain, Ireland and
Scandinavia (but recent record in southernmost
Sweden)
oligolectic on Echium (Boraginaceae)
Hoplitis benoisti (Alfken) southwestern Europe (Iberian peninsula, southern
France, Corsica, Sardinia and possibly Sicily)
oligolectic on Echium (Boraginaceae)
Hoplitis bihamata (Costa) Corsica, Sardinia mesolectic on Echium
(Boraginaceae), Lamiaceae and
Fabaceae
Hoplitis carinata (Stanek) southeastern Europe (Croatia, Macedonia, Bulgaria,
Greece)
oligolectic on Fabaceae with strong
preference for Hedysareae (e.g.
Onobrychis)
Hoplitis fabrei Zanden southwestern Macedonia and Greece excluding
Aegean Islands
polylectic with preference for Echium
(Boraginaceae), Campanulaceae and
Fabaceae
Hoplitis fertoni (Pérez) Spain, Sicily oligolectic on Echium (Boraginaceae)
Hoplitis galichicae spec. nov. Macedonia possibly oligolectic on Sedum
(Crassulaceae)
Hoplitis hilbera Müller western Spain from Cataluna southwards to
Andalucia
mesolectic on Fabaceae, Echium
(Boraginaceae) and possibly
Brassicaceae
Hoplitis holmboei
(Mavromoustakis)
Cyprus oligolectic on Boraginaceae
(Onosma, Echium, Lithodora)
Hoplitis insularis
(Schmiedeknecht)
southwestern Europe (Spain, Balearic Islands,
Sardinia, Sicily)
mesolectic on Echium (Boraginaceae)
and Fabaceae with strong preference
for Echium
Hoplitis jheringii (Ducke) southeastern Europe (northeastern Italy, Slovenia,
Croatia, Macedonia, Greece)
oligolectic on Fabaceae with strong
preference for Loteae
Hoplitis lepeletieri (Pérez) Alps (France, Italy, Switzerland, Germany, Austria,
Slovenia), some low mountain ranges in southern
Germany, Pyrenees (France, Spain) and possibly
Hungary
oligolectic on Echium (Boraginaceae)
Hoplitis lithodorae Müller southwestern Spain to southern France probably oligolectic on Lithodora
(Boraginaceae)
Hoplitis loti (Morawitz) Alps (France, Italy, Switzerland, Germany, Austria,
Slovenia) and Pyrenees (Spain, France)
mesolectic on Fabaceae, Crassulaceae
and Echium (Boraginaceae) with
strong preference for Loteae
Hoplitis manicata (Morice) eastern and southeastern Europe (easternmost
Austria, Czech Republique, Slowakia, Hungary,
Romania, Ukraine, eastern and northeastern Italy,
Croatia, Serbia, Montenegro, Albania, Macedonia,
Greece including Aegean Islands, Bulgaria)
oligolectic on Echium (Boraginaceae)
Hoplitis marchali (Pérez) Iberian Peninsula, Corsica, Sicily oligolectic on Echium (Boraginaceae)
Hoplitis mucida (Dours) southwestern Europe (Iberian peninsula, southern
France, Sicily, northern Italy, southern Switzerland)
oligolectic on Echium (Boraginaceae)
Hoplitis ochraceicornis
(Ferton)
southwestern Europe (Iberian peninsula, Andorra,
French Pyrenees and French Alps, northwestern
Italy)
mesolectic on Echium
(Boraginaceae), Fabaceae and
Lamiaceae
......continued on the next page
MÜLLER
176
·
Zootaxa 4111 (2) © 2016 Magnolia Press
Acknowledgments
Max Schwarz (Ansfelden) provided megachilid material for study. J. van Leeuwen (University of Bern) confirmed
the identity of Sedum pollen in the pollen loads of H. galichicae. V. Matevski (Ss. Cyril and Methodius University,
Skopje) provided information on the occurrence of Crassulaceae species in the Galichica national park. H. Baur
(Natural History Museum Bern) kindly provided access to a digital imaging system for taking photomicrographs.
Comments by two anonymous reviewers were very helpful.
Literature
Cane, J.H. & Sipes, S. (2006) Characterizing floral specialization by bees: analytical methods and a revised lexicon for
oligolecty. In: Waser, N.M. & Ollerton, J. (Eds.), Plant-pollinator interactions from specialization to generalization.
University of Chicago Press, Chicago, pp. 99–122.
Michener, C.D. (2007) The bees of the world. 2
nd
Edition. Johns Hopkins University Press, Baltimore and London, 953 pp.
Müller, A. (2006) Unusual host plant of Hoplitis pici, a bee with hooked bristles on its mouthparts (Hymenoptera:
Megachilidae: Osmiini). European Journal of Entomology, 103, 497–500.
http://dx.doi.org/10.14411/eje.2006.064
Müller, A. (2012) New European bee species of the tribe Osmiini (Hymenoptera: Apoidea: Megachilidae). Zootaxa, 3355,
29–50.
Müller, A. (2015) Palaearctic osmiine bees - systematics and biology of a fascinating group of solitary bees. ETH Zürich.
Available from: http://blogs.ethz.ch/osmiini (accessed 20 November 2015)
Müller, A. & Kuhlmann, M. (2008) Pollen hosts of western palaearctic bees of the genus Colletes (Colletidae) - the Asteraceae
paradox. Biological Journal of the Linnean Society, 95, 719–733.
http://dx.doi.org/10.1111/j.1095-8312.2008.01113.x
Sedivy, C., Dorn, S. & Müller, A. (2013a) Evolution of nesting behaviour and cleptoparasitism in a selected group of osmiine
bees (Megachilidae). Biological Journal of the Linnean Society, 108, 349–360.
http://dx.doi.org/10.1111/j.1095-8312.2012.02024.x
Sedivy, C., Dorn, S., Widmer, A. & Müller, A. (2013b) Host range evolution in a selected group of solitary bees: the
Boraginaceae-Fabaceae paradox. Biological Journal of the Linnean Society, 108, 35–54.
http://dx.doi.org/10.1111/j.1095-8312.2012.02013.x
TABLE 1. (Continued)
Species European distribution area Pollen host preferences
Hoplitis pallicornis (Friese) southeastern Europe except mainland Greece (eastern
and northeastern Italy, Slovenia, Croatia, Albania,
Macedonia, Aegean Islands, Bulgaria)
polylectic with preference for
Boraginaceae (mainly Echium),
Fabaceae and Campanulaceae
Hoplitis perambigua (Peters) Canary Islands (Fuerteventura, Lanzarote, Tenerife) possibly oligolectic on Echium
(Boraginaceae)
Hoplitis pici (Friese) southeastern Europe (Romania, Croatia, Montenegro,
Greece including Aegean Islands)
oligolectic on Muscari
(Hyacinthaceae)
Hoplitis ravouxi (Pérez) southwestern, western, central and eastern Europe
(Iberian Peninsula, France, Italy, Luxembourg,
Belgium, Netherlands, Germany, Switzerland,
Austria, Slovenia, Czech Republique, Slovakia,
Hungary, Romania, Belarus); the species seems to be
absent from southeastern Europe, rendering literature
records from Croatia and Macedonia doubtful
mesolectic on Fabaceae, Echium
(Boraginaceae) and Crassulaceae with
strong preference for Loteae
Hoplitis strymonia Tkalcu Bulgaria possibly oligolectic on Fabaceae (e.g.
Trifolium)
Hoplitis submanicata Zanden Sicily probably oligolectic on Echium
(Boraginaceae)
... Demnach ist die Art als vermutlich oligolektisch einzustufen, die vorwiegend an Fabaceae sammelt, da Lotus corniculatus bevorzugt wird. Müller (2016) beschreibt sie als mesolektisch, wobei Fabaceae, Crassulaceae und Echium (Boraginaceae) als Pollenquellen genutzt werden, Loteae-Arten (z.B. Lotus) aber bevorzugt werden. ...
... O. loti ist eine heute rein alpin verbreitete Art. In den Österreichischen und Schweizer Alpen ist sie weit verbreitet (Schwarz et al., 1999;Müller, 2016;Praz et al., 2023). Wiesbauer (2023) stuft sie in Österreich als selten ein. ...
... Auch in den deutschen Alpen kommt sie vor (Westrich, 2019), ist dort aber aktuell nur sehr lokal verbreitet, ohne dass dabei ein Rückgang erkenntlich ist (Bayerisches Landesamt für Umwelt 2021). O. loti ist auch in den Pyrenäen (Spanien, Frankreich) nachgewiesen (Müller, 2016). ...
Article
Full-text available
Infolge neuer Informationen zu historischen Belegen, die sich im Museum für Naturkunde Berlin fanden, hat der Verfasser keine Zweifel, dass die heute nur alpin verbreitete Art Osmia (Hoplitis) loti um das Jahr 1900 auch in Thüringen und Hessen vorkam. Überlegungen führen zu dem Gedanken, dass O. loti begünstigt durch anthropogene und klimatische Einflüsse erst in den letzten Jahrhunderten in Flusstäler außerhalb der Alpen eingewandert sein könnte. Vermutlich ist die Art dort bereits anfangs des 20. Jahrhunderts ausgestorben, weshalb ihre historische Verbreitung nur durch wenige Belege dokumentiert werden kann. Der Ort, an dem Osmia loti im Rheingau gefunden wurde, wird beschrieben und mit historischen und aktuellen Abbildungen dokumentiert. Um diesen Nachweis zu plausibilisieren, werden Lebensdaten des Sammlers Emil Hanau vorgestellt. Abstract: This publication deals with the historical distribution of the bee species Osmia (Hoplitis) loti Morawitz, 1867 in Germany outside the Alps. As a result of new information on historical specimens found in the Museum für Naturkunde Berlin, the author has no doubt that the species, which today is only distributed in the Alps, also occurred in Thuringia and Hesse around the year 1900. Considerations lead to the idea that O. loti may have migrated into river valleys only in the last few centuries. This spread was possibly favored by anthropogenic and climatic influences. The species probably became extinct there at the beginning of the 20th century, which is why its historical distribution can only be documented by a few records. The site in the Rheingau, where O. loti was found, is described and documented with historical and current illustrations. In order to make the record in Hesse more understandable, the life data of the person Emil Hanau are presented.
... At least 64 Palaearctic species are still undescribed, of which at least 50 species belong to the largest subgenus Hoplitis (Hoplitis) (Aubert et al. 2024;. This subgenus comprises several species groups, the largest of which is the H. adunca species group, one of the taxonomically most challenging osmiine bee taxa due to the high morphological uniformity among its species, especially in the female sex (Müller 2016. The nesting biology of Hoplitis is extremely diverse and encompasses the whole diversity observed across the osmiine bees . ...
... These data suggest a preference for the genus Vicia L. by H. andreasmuelleri, perhaps representing narrow oligolecty on a single plant genus (Müller and Kuhlmann 2008). However, more flower visitation data including microscopical analysis of pollen contained in the female scopae are needed to clarify the species' degree of host plant specialization, although many members of the H. adunca species group exclusively or predominantly exploit Fabaceae for pollen (Müller 2016Sedivy et al. 2012b). Remarks. ...
Article
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Hoplitis (Hoplitis) andreasmuellerisp. nov., a member of the H. adunca species group, is described from the vicinity of Kurush (Samurskiy National Park, Dagestan, Russia). The new species is closely related to H. dagestanica Fateryga, Müller & Proshchalykin, 2023. Females of H. andreasmuelleri can be easily distinguished from those of H. dagestanica by a black antenna, a small impunctate triangular area at the base of the clypeus, ferruginous tarsi, sparse and often interrupted apical band of hairs on terga 1–5, and a whitish scopa. Males of H. andreasmuelleri can be easily distinguished from those of H. dagestanica by a black antenna, the last antennal article not tapering towards the apex, antennal articles 5–12 not modified, ferruginous tarsi, and the lateral lobes of the bilobed membranous appendage at the apical margin of sternum 6 laterally elongated into a rounded tip and medially not separated from each other by an emargination. Females of H. andreasmuelleri were recorded at flowers of two species of the genus Vicia L. (Fabaceae). Hoplitis (Kumobia) abbreviata (Morawitz, 1875) is reported from Russia (Altai Republic) for the first time. New distributional records are reported for H. (Alcidamea) fulva (Eversmann, 1852) and H. (Anthocopa) perezi (Ferton, 1894). Hoplitis (Anthocopa) papaveris (Latreille, 1799) is excluded from the fauna of Crimea and the south of European Russia from where it was previously reported based on a misidentification of H. perezi. The biology of H. perezi in Crimea is briefly discussed. An updated distributional list of all 39 species of Hoplitis known from Russia is provided.
... The subgenus comprises several species groups. Among them, the Hoplitis adunca species group is the largest and one of the taxonomically most challenging osmiine bee taxa due to the high morphological uniformity among its species, especially in the female sex (Müller 2016(Müller , 2023. Members of this species group nest either in various pre-existing cavities (such as rock and stone crevices, hollow stems, abandoned nests of other bees and wasps, insect burrows in wood, rarely empty snail shells) or construct cells freely on the surface of rocks or stones, usually in depressions (Sedivy et al. 2013). ...
... One male of H. dagestanica was also observed on flowers of Astragalus. However, as this was the only flower visiting record for this species, any assumption on its pollen host preference is premature, although many members of the H. adunca species group exclusively or predominantly exploit Fabaceae for pollen (Müller 2016(Müller , 2023Sedivy et al. 2012b). ...
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Hoplitis astragali sp. nov., a member of the H. monstrabilis species group, and H. dagestanica sp. nov., a member of the H. adunca species group, are described. The former species is known from Dagestan in Russia, Azerbaijan, and Turkmenistan, the latter only from Dagestan. Nests of H. astragali are described. Females of this species excavated burrows in a vertical clay cliff, but sometimes chose a horizontal surface for nest excavation, particularly at the entrance of old burrows of Xylocopa olivieri (Apidae). The nest burrows of H. astragali were either sub-vertical or sub-horizontal. The nests were composed of one to three brood cells, an empty vestibule in front of the outermost cell, and a closing plug at the nest entrance made of moistened mud. The inner surface of the cells was covered with a thin wall composed of compact soil, most probably built by the female bee after cell excavation. The pollen loaf was very liquid and had a spherical shape. The egg was deposited on its top. The cocoon consisted of a single thin layer, which uniformly covered the whole inner surface of the cell. There was one generation per year. The prepupae hibernated. Sapyga caucasica (Sapygidae) was recorded in the nests as a kleptoparasite. Both females and males of H. astragali exclusively visited flowers of two species of the genus Astragalus (Fabaceae).
... Pendant longtemps, une seule espèce a été signalée comme hôte : Hoplitis (Hoplitis) heringii (Ducke, 1898) [Matthey, 1949]. Ce Mégachilidae, absent de la faune française [Rasmont et al., 1995 ;Müller, 2016] ...
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... This peculiar form of H. adunca with a short vertex was first mentioned by Müller (2016) but its status remained unclear to date. Both taxa were collected in number by C. Ferton at the same localities and date. ...
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In a recent catalogue of Megachilidae from Corsica, four potentially new species were noted and are described here. Aglaoapis sparsepunctata n. sp., the second European species of the subgenus Aglaoapis Cameron, 1901, is described from the Corsican mountains. Chelostoma (Foveosmia) incisa n. sp. is a species with an intermediate morphology between C. distinctum (Stöckhert, 1929) and C. campanularum (Kirby, 1802). Hoplitis (Alcidamea) agnielae n. sp. is a vicariant species of H. acuticornis (Dufour & Perris, 1840). Hoplitis (Hoplitis) legoffi n. sp. is a member of the adunca complex and appears to be restricted to coastal environments.
... The two additional species groups are the bassana and the monstrabilis groups (Sedivy et al. 2013c). While several new Hoplitis (Hoplitis) species have recently been described from Europe and Morocco (Müller 2012(Müller , 2016(Müller , 2022, more than 50 species belonging to this subgenus, mainly from Turkey, the Levant and Central Asia, remain undescribed (Müller 2023b). ...
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A new osmiine bee species, Hoplitis (Hoplitis) onosmaevaesp. nov. (Megachilidae), is described. So far, this species is exclusively known from the Mercantour National Park in the southwestern French Alps and from mountainous ranges in Turkey and northern Iraq, two areas separated by at least 2000 km. Phylogenetic analyses based on mitochondrial and nuclear genes revealed that H. onosmaevae is closely related to H. adunca (Panzer, 1798), H. benoisti (Alfken, 1935) and H. manicata (Morice, 1901). Hoplitis onosmaevae is presumably narrowly oligolectic and harvests pollen only on flowers of Onosma L. (Boraginaceae). It has a particularly long proboscis, which is probably an adaptation to collect nectar from the long-tubed flowers of this plant genus. The females collect pollen by buzzing the Onosma flowers, a rare behavior in megachilid bees. The species nests in insect burrows in dead wood, similar to H. adunca and H. manicata but unlike other closely related representatives of the subgenus Hoplitis, suggesting a single origin of nesting in dead wood and hollow stems in this lineage. In France, H. onosmaevae inhabits alpine steppe-like habitats close to forests and appears to be extremely local, since only two populations are currently known. The conservation status of this extremely rare bee species in Europe is discussed.
... While Baez & Ortega (1978) listed both above designations, Hohmann & al. (1993) were the first to give species status to Osmia (Hoplitis) perambigua. Michener (2000Michener ( , 2007 and Müller (2016Müller ( , 2018 assign generic rank to Hoplitis and consequently we use Hoplitis (Hoplitis) perambigua (Peters, 1975) for the species treated here. ...
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The construction of nests to rear offspring is restricted to vertebrates and few insect taxa, such as termites, wasps, and bees. Among bees, species of the family Megachilidae are characterized by a particularly high diversity in nest construction behaviour. Many megachilid bees nest in excavated burrows in the ground, others place their brood cells in a variety of above‐ground cavities or attach them to the surface of a substrate, and yet others have adopted a kleptoparasitic habit. Evolutionary transitions between the different nesting sites and between conventional nesting and kleptoparasitism in bees are poorly understood. In the present study, we traced the evolution of nesting site selection and kleptoparasitism in the Annosmia–Hoplitis group (Osmiini), which displays an exceptionally high diversity in nesting behaviour. We found that the evolution of nesting behaviour proceeded unidirectionally from nesting in excavated burrows in the ground to nesting in rock depressions and cavities, followed by the colonization of snail shells and insect borings in dead wood or hollow stems. Kleptoparasitism evolved once and the kleptoparasitic species have derived from the same lineage as their hosts. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 108, 349–360.
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Reviews and defines the range of taxonomic pollen specialization of bees, from monolecty to broad polylecty, with specific parameters for classification. A new category, "mesolecty", is suggested to replace the oxymoron of "narrow polylecty" as well as "eclectic oligolecty" for those species that seem to use pollen from one or a few genera in several unrelated families of flowering plants
Article
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To assess the pollen hosts of 60 western palaearctic bee species of the genus Colletes (Colletidae), we microscopically analysed 1336 pollen loads of collected females. Twenty-six species (43.3%) were found to be specialized at the level of plant family, subfamily or genus. Thirty-four species (56.7%) proved to be pollen generalists to varying degrees, visiting the flowers of up to 15 different plant families. Flowers of the subfamily Asteroideae (Asteraceae) are by far the most important pollen source, contributing 23.6% to the pollen-plant spectrum of the whole bee genus. The high significance of Asteroideae pollen is due to the large number of specialists: 14 Colletes species belonging to four different taxonomic groups harvest pollen exclusively or predominantly on flowers of the Asteroideae. By striking contrast, Asteroideae pollen plays only a marginal role in the diets of the pollen generalists: it was recorded in only 2.7% of the pollen loads and in seven out of the 34 pollen generalists. Among the few generalists exploiting Asteroideae for pollen, three closely related species have ancestors which were possibly specialized on Asteraceae. The pattern of use of Asteroideae pollen by the Colletes bees supports recent findings that this pollen possesses unfavourable or protective properties, which render its digestion difficult, and suggests that bees need physiological adaptations to successfully utilize it. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95, 719–733.
The bees of the world. 2 nd Edition
  • C D Michener
Michener, C.D. (2007) The bees of the world. 2 nd Edition. Johns Hopkins University Press, Baltimore and London, 953 pp.
Ansfelden) provided megachilid material for studyUniversity of Bern) confirmed the identity of Sedum pollen in the pollen loads of H. galichicae
  • Max Schwarz
Max Schwarz (Ansfelden) provided megachilid material for study. J. van Leeuwen (University of Bern) confirmed the identity of Sedum pollen in the pollen loads of H. galichicae. V. Matevski (Ss. Cyril and Methodius University, Skopje) provided information on the occurrence of Crassulaceae species in the Galichica national park. H. Baur (Natural History Museum Bern) kindly provided access to a digital imaging system for taking photomicrographs. Comments by two anonymous reviewers were very helpful.
Palaearctic osmiine bees -systematics and biology of a fascinating group of solitary bees. ETH Zürich Available from
  • A Müller
Müller, A. (2015) Palaearctic osmiine bees -systematics and biology of a fascinating group of solitary bees. ETH Zürich. Available from: http://blogs.ethz.ch/osmiini (accessed 20 November 2015)
Host range evolution in a selected group of solitary bees: the Boraginaceae-Fabaceae paradox (Continued) Species European distribution area Pollen host preferences Hoplitis pallicornis (Friese) southeastern Europe except mainland Greece (eastern and northeastern Italy
  • C Sedivy
  • S Dorn
  • A Widmer
  • A Müller
Sedivy, C., Dorn, S., Widmer, A. & Müller, A. (2013b) Host range evolution in a selected group of solitary bees: the Boraginaceae-Fabaceae paradox. Biological Journal of the Linnean Society, 108, 35–54. http://dx.doi.org/10.1111/j.1095-8312.2012.02013.x TABLE 1. (Continued) Species European distribution area Pollen host preferences Hoplitis pallicornis (Friese) southeastern Europe except mainland Greece (eastern and northeastern Italy, Slovenia, Croatia, Albania, Macedonia, Aegean Islands, Bulgaria)