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Palaearctic Osmia bees of the subgenus Hoplosmia (Megachilidae, Osmiini): Biology, taxonomy and key to species

April 2018
Zootaxa 4415(2):297

April 2018
4415(2):297

DOI:10.11646/zootaxa.4415.2.4

Authors:

ETH Zurich

Hoplosmia, a subgenus of the osmiine bee genus Osmia (Megachilidae), comprises 21 species restricted to the Palaearctic region. Analysis of female pollen loads and field observations indicate that probably all O. (Hoplosmia) species are specialized on Asteraceae except for one pollen generalist species, which exhibits a preference for the pollen of Cistaceae. Among the Asteraceae specialists, differences exist with respect to the three main Asteraceae subfamilies exploited for pollen, with some species exclusively visiting Carduoideae, others exploiting only Asteroideae and Cichorioideae and again others collecting pollen on Asteroideae, Carduoideae and Cichorioideae. All O. (Hoplosmia) species build their brood cells within preexisting cavities: several species exclusively nest in empty snail shells, few species use small cavities in rock and stones and the remaining species colonize linear cavities in dead wood and plant stems or nest in abandoned burrows of other bees and wasps. Chewed leaves serve as material to construct brood cell partitions and nest plug except for two species, which use mud as nest building material. The taxonomic revision of O. (Hoplosmia) revealed the existence of an undescribed species, O. centaureae spec. nov., which occurs in a small area that ranges from the Dead Sea over the Jordan Valley to northernmost Israel. Due to clear morphological gaps and widely disjunct distribution with the nominotypical subspecies, O. pinguis carbo (Zanden 1974) is elevated to species rank. Based on morphology and biology, three species groups are recognized within Hoplosmia. Identification keys for all O. (Hoplosmia) species are given including the hitherto unknown male or female sex of three species.

Flower-visiting females of Osmia ( Hoplosmia). 1: Osmia spinulosa on Buphthalmum salicifolium (Asteroideae). 2: Osmia spinigera on Centaurea spec. (Carduoideae; photo H. Wiesbauer). 3: Osmia bidentata on Centaurea spec. (Carduoideae; photo A. Gogala). 4: Osmia pinguis on Inula spec. (Asteroideae).

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11: Female of Osmia carbo. 12. Female of Osmia pinguis. 13: Male sterna 2-3 of Osmia carbo. 14: Male genitalia of Osmia carbo. 15: Male genitalia of Osmia pinguis.

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Accepted by J. Gibbs: 9 Mar. 2018; published: 30 Apr. 2018

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https://doi.org/10.11646/zootaxa.4415.2.4

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Palaearctic Osmia bees of the subgenus Hoplosmia (Megachilidae, Osmiini):

biology, taxonomy and key to species

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

Hoplosmia, a subgenus of the osmiine bee genus Osmia (Megachilidae), comprises 21 species restricted to the Palaearctic

region. Analysis of female pollen loads and field observations indicate that probably all O. (Hoplosmia) species are spe-

cialized on Asteraceae except for one pollen generalist species, which exhibits a preference for the pollen of Cistaceae.

Among the Asteraceae specialists, differences exist with respect to the three main Asteraceae subfamilies exploited for

pollen, with some species exclusively visiting Carduoideae, others exploiting only Asteroideae and Cichorioideae and

again others collecting pollen on Asteroideae, Carduoideae and Cichorioideae. All O. (Hoplosmia) species build their

brood cells within preexisting cavities: several species exclusively nest in empty snail shells, few species use small cavities

in rock and stones and the remaining species colonize linear cavities in dead wood and plant stems or nest in abandoned

burrows of other bees and wasps. Chewed leaves serve as material to construct brood cell partitions and nest plug except

for two species, which use mud as nest building material. The taxonomic revision of O. (Hoplosmia) revealed the exis-

tence of an undescribed species, O. centaureae spec. nov., which occurs in a small area that ranges from the Dead Sea over

the Jordan Valley to northernmost Israel. Due to clear morphological gaps and widely disjunct distribution with the nom-

inotypical subspecies, O. pinguis carbo (Zanden 1974) is elevated to species rank. Based on morphology and biology,

three species groups are recognized within Hoplosmia. Identification keys for all O. (Hoplosmia) species are given includ-

ing the hitherto unknown male or female sex of three species.

Key words: Apiformes, Asteraceae, Cistaceae, host-plant choice, Hymenoptera, nesting behaviour, oligolecty, polylecty,

snail-shell nesting

Introduction

Hoplosmia Thomson is a subgenus of the osmiine bee genus Osmia Panzer (Megachilidae, Megachilinae, Osmiini)

including 21 species, which are restricted to the Palaearctic region. Its distribution area ranges from Europe and

northern Africa eastwards to Central Asia and eastern Siberia. Hoplosmia was formerly treated at the generic rank

(Michener 2007; Ungricht et al. 2008). However, recent molecular phylogenetic studies revealed that it has derived

from within the genus Osmia (Praz et al. 2008; Rightmyer et al. 2013). Osmia (Hoplosmia) species differ from

most other members of Osmia by their linear rather than punctiform parapsidal lines. Osmia (Hoplosmia) is sister

to a clade containing the subgenera Nasutosmia Griswold & Michener, Allosmia Tkalců and Neosmia Tkalců (Praz

et al. 2008; Rightmyer et al. 2013). Interestingly, all species of that clade, for which the nesting biology is known,

nest in empty snail shells, as many O. (Hoplosmia) species do, suggesting a possibly single origin of snail-shell

nesting at the base of the clade composed of Nasutosmia, Allosmia, Neosmia and Hoplosmia.

After the seminal publication of Tkalců (1974a) on Hoplosmia, which clarified the taxonomy of most species

and contained a number of new species descriptions, Tkalců (1978, 1979, 1992, 1993, 1999) and Zanden (1994)

published several new names, rendering a new revision of O. (Hoplosmia) including updated identification keys

necessary. In the present publication, the subgenus O. (Hoplosmia) is morphologically diagnosed, the current

knowledge on its biology is summarized, the species are revised, one new species is described, and identification

keys are given, which include the hitherto unknown female of O. warnckei (Tkalců 1992) and the unknown males

of O. larochei Tkalců 1993 and O. picena (Tkalců 1999).

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Material and methods

To assess the pollen hosts of the species, scopal pollen contents of all available females were analysed by light

microscopy. The pollen grains were stripped off the scopae with a fine needle and embedded in glycerol gelatin on

a slide. Pollen was identified at a magnification of 400x to family, subfamily, or genus level with the aid of the

literature cited in Westrich & Schmidt (1986), Beug (2004) and an extensive reference collection. Pollen types

represented by less than 5% of the counted grains were excluded to prevent a potential bias caused by

contamination. Morphological terminology and definitions for body measurements follow Michener (2007) with

the following specifications and alterations: i) the distance between lateral ocellus and preoccipital margin was

measured in top view rather than in lateral view; ii) the diameter of the lateral ocellus includes 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 not developed in all individuals; thus, “antennal segments (5)6–13

yellowish-brown” means that segment 5 is sometimes yellowish-brown as segments 6–13. Measurements to the

nearest 0.1 mm or 0.5 mm (for body length) were taken using an ocular micrometer on an Olympus VNT

stereomicroscope. Photomicrographs were taken with the digital microscope Keyence VHX-2000. Depositories of

bee specimens listed in the new records section of the species accounts are not given as a large part of the material

was dispersed after it had been identified by the author.

Morphological diagnosis

Osmia (Hoplosmia) species are small to medium-sized bees (body length 5–11 mm) with linear parapsidal lines

and usually whitish to yellowish tergal hair bands, which are continuous or interrupted. They are the only osmiine

bees, except for some Heriades (Heriades), that have distinctly spined axillae. They differ from these distantly

related Heriades species by the shape of the basal area of the propodeum, which is steeply slanting to vertical

rather than horizontal, lacking a distinct transverse carina along its posterior margin. In further contrast to

Heriades, the female labrum does not bear a preapical tuft of long erect hairs and male tergum 7 is strongly

sclerotized and never hidden under tergum 6. Additional characters of O. (Hoplosmia) females are the form of the

scopal hairs, which are usually wavy in their apical portion, and the extent of the scopa, which laterally often

extends distinctly upwards, covering the lowermost parts of the terga. Additional characters of O. (Hoplosmia)

males are the preapical transverse carina or swelling on tergum 6, which is often nodose, crenulate or dentate, the

shape of tergum 7, which—depending on the species—is pointed, rounded, emarginate or bifid, and the presence of

a spine on sternum 1 in a number of species.

Biology

Pollen hosts. Osmia (Hoplosmia) species have a strong affinity to Asteraceae as pollen hosts (see species accounts

for details). Except for O. fallax Pérez, all species thoroughly analysed were found to be strictly specialized on this

plant family. The other species will most probably also turn out to be Asteraceae oligoleges when more pollen

loads become available for study. Among the Asteraceae specialists, differences exist in the spectrum of

Asteraceae taxa exploited for pollen. Osmia spinulosa (Kirby) is the only species of O. (Hoplosmia) known to

regularly collect pollen on representatives of all three main subfamilies of the Asteraceae, i.e. Asteroideae,

Carduoideae and Cichorioideae (Fig. 1). In contrast, species, such as O. anceyi Pérez, O. bidentata Morawitz, O.

centaureae spec. nov., O. croatica Friese, O. dido Gribodo, O. distinguenda (Tkalců), O. padri (Tkalců), O. picena

(Tkalců) and O. spinigera Latreille, show a clear or exclusive preference for the Carduoideae (Fig. 2, 3). Other

species, such as O. carbo (Zanden), O. ligurica Morawitz, O. pinguis Pérez and O. scutellaris Morawitz, usually

restrict pollen harvesting to the Asteroideae and Cichorioideae (Fig. 4). In all species observed, pollen uptake from

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OSMIINE BEES OF OSMIA (HOPLOSMIA)

the Asteraceae flower heads involves rapid up and down movements of the metasoma against the pollen-bearing

florets. On the Asteroideae, pollen is taken up directly into the metasomal scopa without the aid of the hind legs,

whereas on Carduoideae and Cichorioideae the hind legs are usually involved in directing the florets under the

seesawing metasoma. The only known polylectic O. (Hoplosmia) species is O. fallax, which exhibits a preference

for flowers of the Cistaceae but exploits at least four additional plant families including Asteraceae. Interestingly,

the scopal hairs of O. fallax are straight and apically spatulate rather than wavy and apically button-like as in all

other O. (Hoplosmia) species. Unfortunately, the phylogenetic position of O. fallax within O. (Hoplosmia) is not

known. If future studies show that O. fallax has evolved from within the Hoplosmia clade, its polylectic habit

would be derived supporting the view that many generalist bee species have evolved from oligolectic ancestors and

that they usually have broadened their diet under maintenance of the ancestral host of the clade from which they

originated (Sedivy et al. 2008). Under this evolutionary scenario, the scopal hairs of O. fallax might have lost their

wavy shape either as an adaptation to the need of transporting pollen of different size, shape, ornamentation and

stickiness or since specialized wavy hairs were no longer required to uptake or transport Asteraceae pollen.

FIGURE S 1–4. Flower-visiting females of Osmia (Hoplosmia). 1: Osmia spinulosa on Buphthalmum salicifolium

(Asteroideae). 2: Osmia spinigera on Centaurea spec. (Carduoideae; photo H. Wiesbauer). 3: Osmia bidentata on Centaurea

spec. (Carduoideae; photo A. Gogala). 4: Osmia pinguis on Inula spec. (Asteroideae).

Nesting biology. All Osmia (Hoplosmia) species, for which the nesting biology is known, nest in preexisting

cavities (see species accounts for details). Depending on the species, three types of cavities serve as nesting sites.

First, several species, such as O. cardo, O. croatica, O. fallax, O. pinguis, O. spinigera, O. spinulosa and probably

all the other representatives of the Osmia spinulosa species group, exclusively nest in empty snail shells of small to

medium size (Fig. 5–7). The nests contain 1–4 brood cells separated by one-layered partitions and are sealed with

an additional wall at the shell opening. The vestibule between nest plug and outermost cell partition is never filled

with small particles as in several snail shell nesting Osmia species of the subgenera Allosmia and Neosmia (Müller

2017 and references therein). Interestingly, the outermost cell partition in the nests of O. carbo, O. croatica, O.

pinguis and O. spinulosa is very thick and robust, probably acting as main barrier against nest predators or parasites.

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FIGURE S 5–10. Nesting behaviour of Osmia (Hoplosmia). 5: Female of Osmia spinulosa entering her nest in an empty shell

of Xerolenta obvia. 6: Female of Osmia spinulosa turning her nest in an empty shell of Zebrina detrita in a protected position.

7: Opened nest of Osmia pinguis in an empty snail shell containing three brood cells. 8: Females of Osmia distinguenda at their

nests in small rock cavities (photo G. Pisanty). 9: Opened nest of Osmia ligurica in a preexisting burrow in a dead stem of

Scolymus hispanicus (photo G. Le Goff). 10: Female of Osmia ligurica at the entrance of her nest in a hollow plant stem (photo

H. Wiesbauer).

Osmia (Hoplosmia) species are not known to glue patches of chewed leaves onto the shell surface as many other

snail shell nesting Osmia species of the subgenera Allosmia, Helicosmia and Neosmia do for unknown reasons

(Müller 2017 and references therein). They do not regularly transport their completed nest to a protected place, bury

it into the ground or hide it under plant material. Osmia spinulosa, however, turns its completed nest so that the

shell opening is directed tightly towards the ground (Fig. 6), which may provide some protection against inclement

weather. Similarly, O. spinigera rolls its nest to a shady place if the shell lies on hot sandy ground, probably to

protect the developing larvae from overheating. The selection of shells hidden under stones, as observed in O.

croatica, may also be a strategy to avoid lethal temperatures for the progeny. Second, O. distinguenda and probably

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OSMIINE BEES OF OSMIA (HOPLOSMIA)

its closest relatives, such as O. dido and O. hermonensis (Tkalců), construct the brood cells singly or in small

clusters in small cavities of rock and stones with the rock surface usually forming part of the cell wall (Fig. 8).

Third, the remaining species, i.e. O. anceyi, O. bidentata, O. ligurica, O. padri and O. scutellaris, nest in linear

cavities, such as dead hollow plant stems, insect borings in dead wood or burrows in the pith of plant stems

excavated by other insects (Fig. 9, 10), and more rarely in abandoned nests of aculeate hymenopterans in steep

faces or horizontal ground. Their nests contain up to 11 linearly arranged brood cells, separated from each other by

one-layered partitions.

Osmia (Hoplosmia) species construct cell partitions, nest plug and occasionally also cell walls with chewed

leaves (“leaf pulp”). Exceptions are O. anceyi and O. bidentata, which use mud as nest building material. In O.

bidentata, the entire brood cells including their walls may be constructed from mud possibly in response to a large

diameter or to thin walls of the nesting cavity.

Voltinism and phenology. Species of O. (Hoplosmia) appear to be univoltine. However, species with a rather

extended flight period, such as O. ligurica and O. scutellaris, might have two generations per year. Osmia

spinulosa was found to be parsivoltine with a proportion of the larvae undergoing metamorphosis only after the

second hibernation (Müller 1994). How widespread parsivoltinism is among O. (Hoplosmia) species remains

unclear.

The phenology of O. (Hoplosmia) covers most of the summer half year. Several species appear in mid spring,

such as O. carbo, O. fallax, O. ligurica, O. pinguis and O. scutellaris, whereas others are active mainly in summer,

such as O. anceyi, O. bidentata, O. croatica, O. olgae, O. padri, O. picena and O. spinulosa.

Taxonomy

Michener (2007) treated Hoplosmia as a genus of its own. Based on Tkalců (1974a), he subdivided it into the three

subgenera Hoplosmia s.tr. with Osmia spinulosa as type species, Odontanthocopa Tkalců with O. bidentata as type

species and Paranthocopa Tkalců with O. pinguis as type species. In contrast to Tkalců (1974a), he did not accept

the subgenus Odonterythrosmia Tkalců with O. fallax as type species but synonymized it with Odontanthocopa.

Two recent molecular phylogenetic studies by Praz et al. (2008) and Rightmyer et al. (2013) found Hoplosmia to

have evolved from within the genus Osmia, which indicates that Hoplosmia should be treated as a subgenus of

Osmia, rendering the subgeneric classification of Tkalců (1974a) and Michener (2007) obsolete. In the present

publication, the subgenus Hoplosmia is divided into three morphologically and biologically well-defined species

groups, which only partly overlap with the former subgenera of Tkalců (1974a) and Michener (2007).

Species accounts

Osmia spinulosa species group

The members of this species group are characterized by the morphology of the mesosoma and the nesting site. In

contrast to all other O. (Hoplosmia) species, the scutellum hides the metanotum when seen from above and its

posterior margin reaches the same level as the metanotum or even overhangs it when seen in profile. All species for

which the nesting biology is known nest in empty snail shells, which are never selected as nesting sites by the

members of the other two species groups.

Osmia (Hoplosmia) carbo (Zanden 1994) stat. nov.

Hoplosmia (Paranthocopa) pinguis carbo Zanden 1994: 1116. Type material: Holotype ♀, “8 km O Yeroham” (Israel), private

collection of M. Schwarz (Ansfelden).

Literature records. EGYPT: Friese (1911).

New records. SYRIA: Tadmur oasis, 9.4.1988 (leg. R. Kinzelbach). ISRAEL AND PALESTINE: Haifa

District: Mt. Carmel, En Ayyala, 27.4.2000 (leg. M. Török); West B a n k : Dead Sea N Qumeran Enot Zuqim Res.,

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6.6.1996 (leg. C. Schmid-Egger); Judean Desert, 7 km S Qumeran, 17.3.1997 (leg. J.G. Rozen, M.S. Engel);

Yehuda plain, Geva'ot HaKhurkhar NP, 21.3.2011 (leg. A. Dorchin); Central District: Qadima, 24.2.2009 (leg. A.

Dorchin); Rehovot, 7.3.2009 (leg. A. Dorchin); Netanya Iris Reserve, 4.4.2012 (leg. J.S. Ascher, A.Payne); Judean

Foothills, Lakhish, 16.3.2013 (leg. T. Shapira); Tel Aviv District: Tel Aviv, 3–10.4.1988 (leg. Guichard); Southern

District: 10 km S Beersheva, 1.4.1988 (leg. Guichard); Arad, 6.4.1988 (leg. Guichard); Negev, 4 km SW Sede

Boqer, 7.5.1997 (leg. J.G. Rozen); Arava Valley, Nahal Amazyahu, 30.3.2011 (leg. A. Dorchin); 7 km SE of

Nitzana, 31.3.2012 (leg. J.S.Ascher, A.Payne); Negev Mt., Nahal Nizana, 15 km W Mizpe Ramon, 29.4.2013 (leg.

A. Dorchin, D. Bénon, V. Trunz); JORDAN: 20 km S North Shuna, Tall Al Arbatin, 19.4.1996 (leg. M. Halada); 20

km N Madaba, 1.5.1999 (leg. W. Schläfle); Al Salt, Wadi Shu’ayb, 15.3.2001 (leg. S. Zaitoun); 10 km N Jerash,

20.4.2002 (leg. M. Snizek); Al-Shawbak, 18.4.2007 (leg. C. Praz, C. Sedivy, A. Müller); Wadi Mujib, King’s

Highway, 19.4.2007 (leg. C. Praz, C. Sedivy, A. Müller); Wadi al Hasa S Al Karak, 20.4.2007 (leg. C. Praz, C.

Sedivy, A. Müller).

FIGURES 11–15. 11: Female of Osmia carbo. 12. Female of Osmia pinguis. 13: Male sterna 2–3 of Osmia carbo. 14: Male

genitalia of Osmia carbo. 15: Male genitalia of Osmia pinguis.

Distribution. Levant (Syria, Israel and Palestine, Jordan) and northern Egypt.

Pollen hosts. Oligolectic on Asteraceae (based on 23 pollen loads from 15 different localities in Israel and

Jordan and on field observations). The species exhibits a near exclusive preference for Cichorioideae and

Asteroideae as pollen hosts: 17 pollen loads exclusively consisted of pollen of Cichorioideae (n = 11) or

Asteroideae (n = 6), while six were mixed loads of pollen of both subfamilies. One load contained a substantial

amount of pollen of Carduoideae beside pollen of Cichorioideae and Asteroideae, suggesting that Carduoideae are

occasionally also exploited. Flower records: Andryala spec., Crepis aculeata, C. aspera, Anthemis spec.,

Chrysanthemum coronarium, Senecio vernalis (label records).

Nesting biology. The nests are built in empty snail shells of small to medium size with a diameter of 13–25

mm (Mavromoustakis 1948; A. Müller unpublished data). Twenty-five nests from Jordan contained one (n = 2),

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OSMIINE BEES OF OSMIA (HOPLOSMIA)

two (n = 9) or three (n = 14) brood cells. The brood cells are separated from each other by one-layered partitions

consisting of leaf pulp. An additional wall of leaf pulp seals the shell at or shortly behind its opening. Similar to O.

spinulosa, the nests are not sealed against their rear end with a basal wall and the outermost cell partition is

distinctly more robust than all the other cell partitions and the nest plug, probably acting as a barrier against

parasites and predators. In contrast to O. spinulosa, the shells are never turned after they have been sealed.

Note. Osmia carbo was treated by Zanden (1994) as a subspecies of O. pinguis Pérez 1895. As the distribution

areas of the two taxa are widely separated and the morphological differences are substantial (see Fig. 11–15 and

key to species), O. pinguis carbo is considered here to be a species of its own, i.e. O. carbo (Zanden 1994), stat.

nov.

Osmia (Hoplosmia) centaureae spec. nov.

Holotype. JORDAN: Wadi Mujib, King’s Highway, 19.4.2007, ♂ (leg. C. Praz, C. Sedivy, A. Müller). Deposited

in the Entomological Collection of ETH Zurich.

Paratypes. ISRAEL AND PALESTINE: Northern District: Mt. Tabor, 580 m, 28.5.1991, 3♀, 3♂ (leg. K.

Warncke); 15 km E Qiryat Shemona, Foothill of Hermon, 16.5.1996, 1♀ (leg. O. Niehuis); Lake Tiberias, 2 km

NNE Tiberias, -70 m, 25.5.2011, 2♀, 1♂ (leg. S. Risch). JORDAN: Jordan Valley, Mubalath, 18.4.1996, 8♀, 2♂,

27.4.1996, 9♀, 3♂ (leg. M. Halada); Jordan Valley, 20 km S North Shuna, Tall al Arbatin, 19.4.1996, 2♀, 1♂ (leg.

M. Halada); Jordan Valley, South Shuna, 25–26.4.1996, 1♂ (leg. M. Halada); Jordan Valley, Dayr Alla, 27.4.1996,

3♀ (leg. M. Halada); Jordan Valley, North Shuna, 29–30.4.1996, 6♀, 4♂ (leg. M. Halada); Wadi Mujib, King’s

Highway, 19.4.2007, 10♀, 5♂ (leg. C. Praz, C. Sedivy, A. Müller); Wadi al Hasa, S Al-Karak, 20.4.2007, 2♂ (leg.

C. Praz, C. Sedivy, A. Müller); Wadi Shu’ayb, 20 km W Amman, 22.4.2007, 4♀, 7♂ (leg. C. Praz, C. Sedivy, A.

Müller); Jordan Valley, Tabaqat Fahl, 24.4.2007, 1♂ (leg. C. Praz, C. Sedivy, A. Müller). Deposited in the

Entomological Collection of ETH Zurich, the Oberösterreichische Landesmuseum Linz and the private collections

of M. Schwarz (Ansfelden) and the author.

Other records. ISRAEL AND PALESTINE: Northern District: Dishon, 15.5.1973, 1♀ (leg. H. Bytinski-

Salz). West Bank: Jericho, Wadi Qilt, 21.4.1990, 1♀, 1♂ (leg. K. Warncke).

Diagnosis. Osmia centaureae is in both sexes very similar to O. spinigera (Fig. 16). In contrast to the latter

species, its preoccipital margin is not distinctly raised (Fig. 18), the inner margin of the apicalmost part of the fore

tibial spur is usually slightly convex rather than concave (Fig. 20), the antenna is partly reddish-brown to orange in

its apical half rather than uniformly dark (Fig. 16) and the apical margins of terga 1–2 are often more or less

reddish-brown rather than entirely black. In addition, the female of O. centaureae differs from O. spinigera by the

colour of the scopa and the pilosity of vertex, scutum and scutellum, which is yellowish-white even in fresh

specimens rather than yellowish-red to yellowish-brown (Fig. 16). The male of O. centaureae lacks the dense tuft

of hairs directed alongside the apicalmost part of the gonoforceps typical of O. spinigera (Fig. 24) and the

indistinct and shallow punctation of the shagreened basal part of tergum 6 is almost imperceivable (Fig. 22) and

thus even weaker than in O. spinigera.

Description. FEMALE: Body length 7.5–9 mm. Head: Head as long as wide. Distance between lateral ocellus

and preoccipital margin 2.5–2.75x as long as ocellar diameter. Second segment of labial palpus about 2.5x as long

as first segment. Maximal width of genal area about 0.6x as long as maximal width of compound eye. Mandible

three-toothed. Preoccipital margin sharp but not distinctly raised (Fig. 18). Punctation of clypeus and supraclypeal

area very dense without distinct interspaces; apical third of clypeus with very small punctures, which are about one

third as large as the punctures on the basal two thirds of the clypeus and about half as large as the punctures on the

supraclypeal area. Face covered with white pilosity, which is rather dense and long on frons, paraocular area and

posterior margin of vertex but only sparse and usually short on clypeus, supraclypeal area and vertex. Anterior side

of antennal segments 7–11(12) usually more or less reddish-brown (Fig. 16). Mesosoma: Axilla spined. Posterior

margin of scutellum flattened, reaching in profile well beyond metanotum. Punctation of scutum and scutellum

very dense with interspaces usually not exceeding the diameter of half a puncture except for the lateral part of the

scutum where the punctation is often more scattered. Punctation of mesepisternum very dense without distinct

interspaces. Scutum moderately densely and scutellum very densely haired, longest hairs on scutum about 5x as

long as the diameter of one puncture. Pilosity of mesepisternum long and rather dense, partly hiding the sculpture

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of the integument. Pilosity of scutum and scutellum yellowish-white, of mesepisternum white (Fig. 16). Tegula

dark reddish-brown to black. Inner margin of apicalmost part of fore tibial spur slightly convex (Fig. 20).

Metasoma: Punctation of terga 1–3 very dense with interspaces rarely reaching the diameter of one puncture, of

terga 4–6 even denser with interspaces not exceeding the diameter of half a puncture. Apical margins of terga

impunctate and often more or less reddish-brown. Terga 1–6 with white apical hair bands, which are medially

interrupted on tergum 1 and often also on tergum 2 (Fig. 16). Scopa yellowish-white (Fig. 16).

MALE: Body length 7–9.5 mm. Head: Head about 0.9x long as wide. Distance between lateral ocellus and

preoccipital margin 2.6–2.9x as long as ocellar diameter. Second segment of labial palpus about 2.6x as long as

first segment. Maximal width of genal area about 0.5x as long as maximal width of compound eye. Mandible two-

toothed. Preoccipital margin sharp but not distinctly raised. Frons, paraocular area, supraclypeal area and clypeus

covered with dense white pilosity. Antennal segments (5)6–13 more or less yellowish-brown to orange. Mesosoma:

Axilla spined. Posterior margin of scutellum flattened, reaching in profile well beyond metanotum. Punctation of

scutum, scutellum and mesepisternum as in the female. Scutum and scutellum covered with dense greyish-white

pilosity. Pilosity of mesepisternum long, dense and white. Tegula dark reddish-brown to black. Inner margin of

apicalmost part of fore tibial spur straight to slightly convex. Metasoma: Punctation of terga 1–5 very dense with

interspaces rarely reaching the diameter of one puncture. Tergum 6 basally densely shagreened and with very weak

and almost imperceivable punctation (Fig. 22). Preapical margin of tergum 6 with long spines (Fig. 22). Tergum 7

with single median tooth (Fig. 22). Apical margins of terga 1-5 impunctate and usually more or less reddish-brown,

often very weakly bent upwards. Terga 1–5 with white apical hair bands, which are medially interrupted on tergum

1 and often also on tergum 2. Sternum 1 with long bifurcated spine (Fig. 17). Sternum 2 with preapical transverse

swelling (Fig. 17). Apical margin of sternum 4 medially very slightly emarginated (Fig. 17). Apical margin of

sternum 5 medially with wide and rather deep emargination, which is densely beset with yellowish hairs (Fig. 17).

Sternum 6 distinctly concave, its apical margin evenly rounded (Fig.17). Punctation of sterna 2–4 moderately

dense, interspaces only rarely exceeding the diameter of two punctures (Fig. 17). Apical margins of sterna 2-4 beset

with whitish hairs, which are laterally longer and denser than medially (Fig. 17). Apical third of gonoforceps with

numerous erect white hairs (Fig. 24).

Distribution. Known only from a restricted area, which extends from the southeastern border of the Dead Sea

in Jordan over the Jordan Valley to northernmost Israel. At five localities in Israel and Jordan, O. centaureae was

collected together with O. spinigera on the same day indicating syntopic occurrence of these two closely related

species within the distribution range of O. centaureae.

Pollen hosts. Oligolectic on Asteraceae (based on 28 pollen loads from 9 different localities in Israel and

Jordan and on field observations). Twenty-six pollen loads were pure loads of Centaurea pollen, while the other

two were mixed loads of Centaurea and thistle pollen, suggesting that O. centaureae probably restricts pollen

harvesting to the subfamily Carduoideae.

Nesting biology. Unknown.

Etymology: The specific name refers to the species’ most important pollen host, Centaurea (Asteraceae,

Carduoideae).

Osmia (Hoplosmia) croatica Friese 1893

Osmia croatica Friese 1893: 353. Type material: Lectotype ♂, by designation of Tkalců (1974a), “Triest” (Italy), Museum für

Naturkunde Berlin.

Literature records. ITALY: Triest (Tkalců 1974a); Friuli-Venezia Giulia, Toscana, Umbria (Pagliano 1994).

SLOVENIA: Submediterranean region (Gogala 2014). CROATIA: Ugljan, Kali, Krk/Baska (Tkalců 1974a);

Karlobag (Józan 2009). SERBIA: Ungricht et al. (2008). GREECE: Portaria/Pelionpass, Mykene (Zanden 1983);

Lesbos (Grace 2010). BULGARIA: Slantschev Brjag (Tkalců 1974a); Koneso, Varna (Zanden 1983). TURKEY:

Eskisehiri/Inönü, 20 km S Ankara (Zanden 1994); Silivri/Istanbul (Zanden 1989).

New records. ITALIA: Toscana: 25 km SSW Volterra, Canneto, 13–25.7.1997 (leg. S. Risch); Abruzzo: Forme

near Avezzano, 1000m, 27.8.2002, 31.8.2007 (leg. A. Müller); Molise: Gurdialfiera, 10.6.2007 (leg. G. Pagliano);

Puglia: Mte. Gargano, San Marco in Lamis, Macchione, 500 m, 2.7.1994 (leg. M. Bernasconi, A. Müller); Lesina,

17.6.2007 (leg. G. Pagliano). CROATIA: Istria: 10 km N Pula, 25–27.8.1998 (leg. J. Halada); 10km E Rovinj,

305

OSMIINE BEES OF OSMIA (HOPLOSMIA)

16.7.2002 (leg. Z. Pedr); Vodnjan, 18.7.2002 (leg. M. Halada); Laborika, 20.7.2002 (leg. M. Halada); Umag, 5–

12.7.2009 (leg. P. Pacholatko); Krk, S Stara Baska, 17.7.2014 (leg. C. Schmid-Egger); Split, 2.8.1958 (leg. W.

Schläfle); Baska Voda, 7.1972 (leg. K. Polacek); Senj, 13.7.1993 (leg. M. Halada); Makarska, 20.7.2001, 27.6.2011

(leg. J. Linda, Z. Pedr). MONTENEGRO: 10 km W Cetinje, 700 m, 4.9.2014 (leg. J. Halada). GREECE: Ionian

Islands: Kefalonia, Mihalitsata, 7.2009 (leg. P. Banar); Central Macedonia: Litochoro, Plaka, 21–28.7.1988 (leg. S.

Risch); Central Greece: 40 km N Lamia, Domokos, 1.7.1996 (leg. M. Halada); Western Greece: Michas-

Erymanthos mountains, 1300–1700 m, 10.7.1996 (leg. W. Arens); Olympia, Alfios valley, 24.7.1997 (leg. W.

Arens); Andritsena, Vassae, 15.7.2007 (leg. W. Arens); Peloponnese: Kap Tenaro, Mani, 7.6.1996, 29.6.1997 (leg.

W. Arens); Kotili, Likeos Oros, 1000–1400 m, 22.6.1996, 20.7.1997 (leg. P. Hartmann, W. Arens); Kiparissia,

Peristeria, 30.6.1996 (leg. W. Arens); 20 km N Pilos Marathopoli, 8.7.1996 (leg. M. Halada); Alifira, Arkadia,

21.6.1998 (leg. W. Arens); Stymphalia, 8.7.2001 (leg. W. Arens); Aegean Islands: Limnos, Pyrgoi Physinis,

11.6.2012 (leg. A. Chroni). BULGARIA: Varna, Zlatni Pjasecy, 15.7.1957 (leg. Boucek); Slantschev Brjag,

23.7.1968 (leg. M. Kocourek); Kavarna, 5.7.1976 (leg. B. Tkalců); Primorsko, 26.7.1979 (leg. A. Hoffer); Sliven

env., 21.7.1987 (leg. V. Bartak); 30 km W Sofia, 12.8.1993 (leg. M. Halada); Plovdiv, 20.7.1996 (leg. Z. Pedr);

Stara Zagora, 5.7.2000 (leg. M. Snizek). TURKEY: Istanbul: Sile, 1.8.1965 (leg. K. Warncke); Kütahya: 30 km N

Kütahya, 13.6.2000 (leg. M. Halada); Isparta: Karakus Dagi centr., 1460 m, 11.7.2006 (leg. J. Halada); Ankara: 20

km S Ankara, 2.8.1979 (leg. K. Warncke); Konya: Aksehir, 2.8.1991 (leg. K. Warncke); Nevsehir: Nevsehir,

9.8.1972 (leg. K. Warncke); Nigde: Camardi, 1800 m, 10.8.1991 (leg. K. Warncke); Malatya: 5 km E Malatya,

27.6.2000 (leg. M. Halada); Erzincan: 20 km E Tercan, 22.8.1991 (leg. K. Warncke); Artvin: Demirkent/Salekör,

1600 m, 1.9.1995 (leg. P. Hartmann); Bitlis: E Ahlat, 17.8.1991 (leg. K. Warncke); Van : E of Muradiye, 3.7.2000

(leg. M. Halada).

Distribution. From Italy and Slovenia over the Balkan Peninsula (Croatia, Serbia, Montenegro, Greece,

Bulgaria) to easternmost Turkey. The westernmost record is from Volterra in the Italian Toscana province.

Pollen hosts. Oligolectic on Asteraceae (based on 30 pollen loads from 14 different localities in Italy, Croatia

and Greece, on the pollen content of one brood cell and on field observations). As all pollen loads analysed were

pure loads of Centaurea (n = 29) or thistle pollen (n = 1) and the cell provision only contained Centaurea pollen, O.

croatica is likely a specialist of Carduoideae.

Nesting biology. The nests are built in empty snail shells (A. Müller unpublished data). The only nest

discovered contained a single brood cell and was hidden below stones. The nest architecture was found to be the

same as in O. spinulosa as was the material used to construct cell partitions and the nest plug (see species account

of O. spinulosa). In one case, a female was observed to collect leaf pulp from leaves of Scabiosa.

Note. Osmia microgramma Dours 1873 was considered a potential junior synonym of O. croatica by Tkalců

(1974a). As the type material has been destroyed and the original description does not allow one to unambiguously

identify the species, O. microgramma was considered a nomen dubium by Ungricht et al. (2008).

Osmia (Hoplosmia) elegans (Tkalců 1992)

Hoplosmia (Hoplosmia) elegans Tkalců 1992: 220. Type material: Holotype ♂, “Side, 8–20.6.1985” (Turkey), Naturalis

Biodiversity Center Leiden.

Literature records. TURKEY: Mugla: Torba/Budrum(Tkalců 1992).

New records. GREECE: Aegean Islands: Lesbos, 8.9 km SSE Mytilene, 60 m, 27.6.2004 (leg. T. Petanidou);

Lesbos, Eresos, 6.6.2012 (leg. G. Nakas). TURKEY: Aydin: Kusadasi, 27–29.6.2006 (leg. E. Scheuchl); Aydin,

Adnan Menderes University Campus, 11.6.2006, 4.7.2006 (leg. E. Scheuchl); Denizli: 35 km SSE Denizli, 970 m,

5.7.2006 (leg. J. Halada); Mersin: 4 km W Erdemli, 3.7.1995, 220 m (leg. P. Rasmont); Adana: Cicekli, 25 km N

Adana, 50m, 3–5.7.1998 (leg. J. Bezdek). JORDAN: 30 km WWN Ajloun, 30.4.2006 (leg. F. Kantner).

Distribution. From Lesbos in the Aegean Sea eastwards to central Turkey and from Turkey southwards to

northern Jordan.

Pollen hosts. The only pollen load available consisted of pollen of Centaurea. Flower record: Centaurea

solstitialis (label record).

Nesting biology. Unknown.

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FIGURE S 16–25. 16: Female of Osmia centaureae. 17: Male sterna of Osmia centaurea. 18: Female head of Osmia

centaureae in lateral view (pilosity on vertex removed). 19: Female head of Osmia spinigera in lateral view (pilosity on vertex

removed). 20: Female fore tibial spur of Osmia centaureae. 21: Female fore tibial spur of Osmia spinigera. 22: Male tergum 6

of Osmia centaureae. 23: Male tergum 6 of Osmia spinigera. 24: Male genitalia of Osmia centaureae. 25: Male genitalia of

Osmia spinigera.

Osmia (Hoplosmia) fallax Pérez 1895

Osmia fallax Pérez 1895: 13. Type material: Lectotype ♂, by designation of Tkalců (1974a), “Andalusia” (Spain), Muséum

National d’Histoire Naturelle Paris.

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OSMIINE BEES OF OSMIA (HOPLOSMIA)

Literature records. SPAIN: Granada: Sierra Elvira (Moreno-Rueda et al. 2008); Malaga (Tkalců 1974a);

Barcelona, Cataluña (Ceballos 1956).

New records. SPAIN: Va l en c i a : Almenara la Llosa, 12 km N Sagunt, 27.4.2001 (leg. E. Scheuchl); 80 km SW

Valencia, Muela de Cortes reserv., 14.5.2003 (leg. M. Halada); Alicante: Xixona, 1.5.2000 (leg. E. Scheuchl); Coll

de Rates, between Callosa d’en Sarria and Parcent, 520 m, 3.5.2000 (leg. E. Scheuchl); Murcia: Alahama de

Murcia, Monte Muela, 29.4.2000 (leg. E. Scheuchl); Pto. de Jumilla, 800 m, 19.5.2003 (leg. M. Halada); 25 km

SW Cartagena, 19.5.2003 (leg. M. Halada); Almeria: 50 km W Almeria, Berja, 21.4.2003 (leg. M. Halada);

Granada: Polopos, 5.5.1997 (leg. M. Halada). MOROCCO: Fès-Meknès: Ifrane, 15.6.1962 (leg. W. Schläfle);

Ifkern, 25 km E Boulemane, 24–25.5.1995 (leg. M. Halada); Ifrane, 1680 m, 10.6.1996 (leg. P. Rasmont);

TUNISIA: Kasserine: 10 km NNW Thelepte, 24.3.2001 (leg. C. Saure).

Distribution. Western and southern Spain, Maghreb (Morocco, Tunisia).

Pollen hosts. Polylectic with a strong preference for Cistaceae (e.g. Helianthemum), additional pollen sources

include Brassicaceae, Asteraceae (Asteroideae, Cichorioideae), Echium (Boraginaceae) and Lamiaceae (based on

13 pollen loads from six different localities in Spain and Morocco). Pollen of Cistaceae constituted 93.6% of the

total pollen grain volume and was recorded in all pollen loads, seven of which were pure Cistaceae pollen loads,

indicating a very high importance of the flowers of Cistaceae as host plants. Interestingly, the scopal hairs of O.

fallax are straight and apically spatulate rather than wavy and apically button-like as in all the other O. (Hoplosmia)

species, which are most probably all specialized on Asteraceae. This deviating scopal hair shape might be an

adaptation to the need of transporting pollen of different sizes and morphologies or might reflect the relaxed

selection for wavy hairs to uptake or transport Asteraceae pollen. Furthermore, in both sexes of O. fallax the first

and the second segment of the labial palpus are of roughly the same length, which is in sharp contrast to all the

other O. (Hoplosmia) species, where the first segment is distinctly shorter than the second. This difference might be

due to different morphological requirements to suck nectar from the host plants of O. fallax and Asteraceae,

respectively.

Nesting biology. The nests are built in empty snail shells (e.g. Sphincterochila candidissima) and contain 1–4

brood cells per shell (Moreno-Rueda et al. 2008; J. Ortiz-Sanchez personal communication). In southeastern Spain,

O. fallax and the cleptoparasitic megachilid bee species Stelis odontopyga Noskiewicz were found at the same

localities inhabiting empty snail shells (Moreno-Rueda et al. 2008), suggesting that S. odontopyga, which is known

to develop in nests of O. spinulosa (Noskiewicz 1925; Blüthgen 1926), might also parasitize O. fallax.

Osmia (Hoplosmia) picena (Tkalců 1999)

Hoplosmia (Hoplosmia) picena Tkalců 1999: 11. Type material: Holotype ♀, “P.N. Sibillini, 5.8.1996” (Italy), Naturalis

Biodiversity Center Leiden.

Diagnosis of the hitherto unknown ♂: The male of O. picena is characterized by an emarginate spine on sternum

1. The only other O. (Hoplosmia) species with this character are O. centaureae and O. spinigera, which differ from

O. picena by the distinctly coarser and denser punctation of terga and sterna and by the longer ocellooccipital

distance (see Key to species).

New records. ITALY: Abruzzo: Rocca di Mezzo, 1500 m, 21.8.2002 (leg. A. Müller); Rovere near Rocca di

Mezzo, 1350–1400 m, 25.8.2002, 2.8.2010 (leg. A. Müller).

Distribution. Osmia picena has a very restricted distribution and is known only from the Sibillini and Abruzzo

mountains in the Central Apennines of Italy. At three localities in the Abruzzo mountains, O. picena was observed

to fly together with O. spinulosa indicating syntopic occurrence of these two closely related species within the

distribution range of O. picena.

Pollen hosts. Oligolectic on Asteraceae (based on 21 pollen loads from five different localities in the Abruzzo

mountains and on field observations). Eighteen pollen loads exclusively consisted of pollen of Carduoideae

(Centaurea and/or thistles), while the other three loads additionally contained pollen of Cichorioideae (n = 2) and

Asteroideae (n = 1), suggesting that O. picena exhibits a preference for Carduoideae as pollen hosts, but

occasionally also exploits flowers of Cichorioideae and Asteroideae.

Nesting biology. Unknown.

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308

Osmia (Hoplosmia) pinguis Pérez 1895

Osmia pinguis Pérez 1895: 12. Type material: Lectotype ♀, by designation of Tkalců (1974a), “La Calle” (Algeria), Muséum

National d’Histoire Naturelle Paris.

Osmia indivisa Benoist 1928: 214. Type material: Holotype ♂, “Marrakech” (Morocco), Muséum National d’Histoire Naturelle

Paris. Synonymy in Zanden (1985).

Literature records. SPAIN: Ornosa et al. (2006).

New records. MOROCCO: Fès-Meknès: Ifkern, 25 km E Boulemane, 24–25.5.1995 (leg. M. Halada);

Oriental: Figuig, Jbel el Klakh, 30.5.1996 (leg. M. Terzo); Casablanca-Settat: El Jadida, 5.1964 (leg. W. Schläfle);

Marrakesch-Safi: Sidi-Moktar, 17.3.1992 (leg. H.J. Flügel); Drâa-Tafilalet: 700m S Tagounite, 17.4.2014 (leg. R.

Prosi); Souss-Massa: 31 km SE Ait Baha, 4.3.2003 (leg. Ø. Berg); 20 km N Tafraoute, 14.4.2017 (leg. A. Müller);

33 km SW Sidi-Ifni, 18.4.2017 (leg. A. Müller). ALGERIA: M’Sila: 18 km E Bow-Saada, 25.4.1986 (leg.

Gerstmeier); Guelma: Guelma, Campus, 4.5.2008 (leg. S. Aguib); Mila: Thleghma, 5.5.2008 (leg. S. Aguib);

Tebessa: Hammamet, 26.4.2002 (leg. N. Benarfa). TUNISIA: Bizerta: 20 km W Mateur, 30.4.2012 (leg. C. Sedivy,

A. Müller); Nabeul: 5 km NW Grombalia, 11.4.2000 (leg. P. Hartmann); Kef: 2 km SW El Kef, 28.4.2012 (leg. C.

Sedivy, A. Müller); Kairouan: 48 km S Kairouan, 9.4.1994 (leg. M. Schwarz); Kasserine: 10 km NW Thelepte,

27.4.2012 (leg. C. Sedivy, A. Müller); Sidi Bouzid: Sidi Bouzid, 12.4.1999 (leg. K. Denes); Gafsa: Gafsa,

14.4.2001 (leg. M. Halada); Toz e u r : Tamerza, 23.3.2001 (leg. C. Schmid-Egger); Gabes: 20 km N Metlaoui,

26.4.2012 (leg. C. Sedivy, A. Müller); Medenine: Djerba, 3.3.2008 (leg. H. Schwenninger); 10 km S Zarzis,

22.4.2012 (leg. C. Sedivy, A. Müller); Tataouine: 56 km S Tataouine, 11.4.1994 (leg. M. Schwarz).

Distribution. Southernmost Spain and Maghreb (Morocco, Algeria, Tunisia).

Pollen hosts. Oligolectic on Asteraceae (based on 19 pollen loads from 14 different localities in Morocco,

Algeria and Tunisia and on field observations). The species exhibits a near exclusive preference for Cichorioideae

and Asteroideae as pollen hosts (Fig. 4): 15 pollen loads exclusively consisted of pollen of Cichorioideae (n = 11)

or Asteroideae (n = 4), while four were mixed loads of pollen of both subfamilies. Flower records: Chrysanthemum

coronarium, Leontodon spec., Pallenis spinosa (label records).

Nesting biology. The nests are built in empty snail shells of small to medium size with a diameter of 15–30

mm, e.g. Otala lactea (Helicidae); nests were also found in elongate conical shells of Rumina sp. (Subulinidae)

with a length of 24–32 mm and a diameter of 9–12 mm (Ferton 1920, A. Müller unpublished data). Eighteen nests

from Morocco (n = 12) and Tunisia (n = 6) contained two (n = 6), three (n = 9) or four (n = 3) brood cells (Fig. 7).

The brood cells are separated from each other by one-layered partitions consisting of leaf pulp. An additional wall

of leaf pulp seals the shell at or few millimetres behind its opening. The empty vestibule between the outermost cell

and the nest plug varies in length from one fourth to five sixth of one shell whorl. As in O. spinulosa, the nests are

not sealed against their rear end with a basal wall and the outermost cell partition is distinctly more robust than all

the other cell partitions and the nest plug, probably acting as a barrier against parasites and predators. In contrast to

O. spinulosa, the shells are never turned after they have been sealed.

Osmia (Hoplosmia) spinigera Latreille 1811

Osmia spinigera Latreille 1811: 584. Type material: Holotype ♂, “Égypte” (Egypt), type depository unknown (Tkalců, 1974a).

Osmia clavicula Gerstaecker 1869: 347. Type material: Lectotype #m, by designation of Tkalců (1974a), “Naxos” (Greece),

Museum für Naturkunde Berlin. Synonymy in Tkalců (1974a).

Literature records. MACEDONIA: A. Gogala (personal communication). GREECE, Aegean Islands: Rhodos

(Mavromoustakis 1959). TURKEY: Erzincan: Kemaliye (Özbek & Zanden 1992). LEBANON: Tkalců (1974a).

New records. GREECE: Central Macedonia: Litochoro, Plaka, 10–28.7.1988 (leg. S. Risch); Attica: Sounion,

3.6.1963, 15.6.1966 (leg. W. Schläfle); Western Greece: Patras, 11.6.1963 (leg. W. Schläfle); Olympia, Alfios

valley, 5.7.1996 (leg. W. Arens); Neochori, 6 km S Zacharo, 18.6.1997 (P. Hartmann); Peloponnese: Methoni,

Castro, 9.5.1995 (leg. W. Arens); Sparta, Amyklai, 19.5.1995 (leg. W. Arens); Examilia, Korinth, 14.6.1995 (leg.

W. Arens); Stymphalia, 15.6.1995 (leg. W. Arens); Kap Tenaro, Mani, 7.6.1996 (leg. W. Arens); Voidokilia near

Pylos, 30.6.1996 (leg. W. Arens); 40 km N Tripoli, Scotini, 3.7.1996 (leg. M. Halada); 20 km W Kalamata,

Peialidi, 6.7.1996 (leg. M. Halada); Mantinea, 7.7.2000 (leg. W. Arens); Ghialova, 5.5.2007 (leg. J. Smit); Aegean

309

OSMIINE BEES OF OSMIA (HOPLOSMIA)

Islands: Chios, Managros, 16.5.2012 (leg. P. Toutziarakis); Kea, Poisses-Koundouros, 27.6.2013 (leg. T.

Petanidou); Lesbos, Sigri, 29.5.2005 (leg. A. Grace); Limnos, Phalakros, 12.6.2012 (leg. A. Chroni, T. Tscheulin);

Naxos, Sagri, 13.6.2012 (leg. I. Vavitsas). BULGARIA: Stara Zagora, 21.6.1963 (leg. Z. Pedr); Slantschev Brjag,,

10.6.1972 (leg. M. Kocourek); Albena, 14.7.1976 (leg. B. Tkalců); Kardzali, Balabanovo, 350 m, 22.6.2007 (leg.

M. Halada). TURKEY: Istanbul: Istanbul, 29.7.1965 (leg. K. Warncke); Mugla: Knidos, 18.4.1981 (leg. K.

Warncke); Baffa lake, 19.4.1996 (leg. P. Hartmann); 4 km ENE Ortaca, 400 m, 25.5.2000 (leg. J. Smit); Eskisehir:

Inönü, 1.8.1991 (leg. K. Warncke); Isparta: Karakus Dagi centr., 1460 m, 11.7.2006 (leg. M. Halada); Antalya:

Side, 8–20.6.1985 (leg. N. Mohr); Ankara: Ankara, 16.6.2006 (leg. E. Scheuchl); Mersin: Kiskalesi bei Silifke, 9–

13.5.1988 (leg. N. Mohr); between Sarikaya and Sarikayak, 920 m, 22.5.2005 (leg. E. Scheuchl); Limonlu,

23.5.2005 (leg. E. Scheuchl); Erzurum: Ispir, 28.6.2008 (leg. J. Rozen). SYRIA: Tartus, 25.5.1996 (leg. M.

Halada). ISRAEL AND PALESTINE: Northern District: Golan, 2 km NW Hamat Gader, 3.5.2010 (leg. C. Sedivy,

C. Praz); Upper Galilee, Nahal Ga´ton, 2 km N Ga´ton, 240m, 20.5.2011 (leg. S. Risch); Lake Tiberias, 2 km NNE

Tiberias, -70 m, 25.5.2011 (leg. S. Risch); Jordan Valley, Tel Qazir, -158 m, 26.5.2011 (leg. A. Dorchin); Lower

Galilee, W Moreshet, 250 m, 28.5.2011 (leg. S. Risch); W Moreshet, 255 m, 28.5.2011 (leg. A. Dorchin); Lower

Galilee, 1.3 km N Tiv'on, 209m, 7.6.2011 (leg. A. Dorchin); Hermon, Majdal Shams, 9.7.2011 (leg. G. Pisanty).

Haifa District: 22 km S Haifa, Nasholim, 18.5.1996 (leg. M. Hauser); Northern Coastal Plain, Ma'agan Mikhael, 7

m, 4.6.2011 (leg. A. Dorchin); Central District: Nes Tsiyona, 31.3.2009 (leg. A. Dorchin); Beit Hanan, 26.4.2009

(leg. A. Dorchin); Tel Yizhaq NR, 20 m, 13.4.2010 (leg. A. Dorchin); 1.75 km SW of Gan Soreq, 18 m, 6.5.2010

(leg. A. Dorchin); Tel Aviv District: Tel Aviv, 3–10.4.1988 (leg. Guichard); Tel Yizhaq, 20 m, 21.5.2011 (leg. A.

Dorchin); Jerusalem District: Judean Foothills, Neve Shalom, 14.5.2009 (leg. G. Bartana); Judean Foothills, Tal

Shahar, 26.4.2010 (leg. L. Gal); Jerusalem, Emek HaArazim, 1.5.2010 (leg. C. Sedivy, C. Praz); West Bank:

Geva'ot, HaKhurkhar NP, 50 m, 1.4.2010 (leg. A. Dorchin); Yehuda mountains, En Perat, 630 m E Anatot,

1.5.2013 (leg. A. Dorchin, D. Bénon, V. Trunz); Southern District: Iris Yeroham NR, 555 m, 28.3.2010 (leg. A.

Dorchin); Negev Mountains, 16 km SE Mizpe Ramon, 815 m, 3.5.2011 (leg. A. Dorchin); Negev Mt., Nahal

Nizana, 10 km WSW Mizpe Ramon, 29.4.2013 (leg. A. Dorchin, D. Bénon, V. Trunz). JORDAN: Jordan Valley,

Mubalath, 18.4–27.4.1996 (leg. M. Halada); Jordan Valley, 20 km S North Shuna, Tall Al Arbatin, 19.4.1996 (leg.

M. Halada); Jordan Valley, South Shuna, 25–26.4.1996 (leg. M. Halada); Jordan Valley, Dayr Alla, 27.4.1996 (leg.

M. Halada); 10 km N Jerash, 20.4.2002 (leg. M. Snizek); Al-Shawbak, 18.4.2007 (leg. C. Praz, C. Sedivy, A.

Müller); Wadi al Hasa S of Al-Karak, 20.4.2007 (leg. C. Praz, C. Sedivy, A. Müller); Jordan Valley, Tabaqat Fahl,

24.4.2007, 1 m (leg. C. Praz, C. Sedivy, A. Müller); 20 km SW Madaba, 400m, 26.5.2007 (leg. Z. Kejval).

Distribution. From the Balkan Peninsula (Macedonia, Greece, Bulgaria) eastwards to eastern Turkey and from

Turkey southwards to the Levant (Syria, Lebanon, Israel and Palestine, Jordan) and northern Egypt.

Pollen hosts. Oligolectic on Asteraceae (based on 29 pollen loads from 23 different localities in Greece, Israel,

Jordan and Syria and on field observations). Twenty-seven pollen loads exclusively consisted of pollen of

Carduoideae (Centaurea and/or thistles), while the other two loads also contained pollen of Asteroideae (n = 1) and

Cichorioideae (n = 1), suggesting that O. spinigera exhibits a strong preference for Carduoideae as pollen hosts

(Fig. 2), but occasionally also exploits flowers of Asteroideae and Cichorioideae. Flower records: Pallenis spinosa

(Mavromoustakis 1959), Centaurea crocodylium, C. hyalolepis, C. iberica, C. pallescens, C. procurrens, C.

spinosa, Glebionis coronaria, Silybum marianum (label records).

Nesting biology. The nests are built in empty snail shells of medium size, e.g. Theba pisana (H. Wiesbauer

personal communication). In sand dune areas in Western Greece near Patras, where the temperature of the soil

surface reaches 50–60°C on sunny days, the females preferentially nested in shells lying in the shade of small

shrubs; if the shells lay openly on the hot sandy ground, the females rolled their nests to sun-protected places under

shrubs, probably to protect them from overheating (H. Wiesbauer personal communication). Cell partitions and

nest plug are constructed from leaf pulp.

Osmia (Hoplosmia) spinulosa (Kirby 1802)

Apis spinulosa Kirby 1802: 261. Type material: Lectotype ♂, by designation of Tkalců (1974a), (United Kingdom), Natural

History Museum London. Type species of Hoplosmia Thomson.

Osmia euchreiformis Radoszkowski 1882: 77. Type material: Holotype ♂, “Echmiadzin” (Armenia), Museum für Naturkunde

Berlin. Synonymy in Tkalců (1974a).

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Literature records. SPAIN: Girona (Ceballos 1956). ANDORRA: Andorra-la-Vella (Zanden 1958). ITALY:

Pagliano (1994). CROATIA: Józan (2009). MACEDONIA: Gorica near Ohrid (Zanden 1984). ROMANIA: Ban-

Calefariu (2009). TURKEY: Aydin, Agri (Özbek & Zanden 1992). UNITED KINGDOM: southern England (Falk

& Lewington 2015). FRANCE: Benoist (1931). BELGIUM: Pauly (1999). NETHERLANDS: Peeters et al.

(2012). LUXEMBOURG: Rasmont et al. (1995). GERMANY: Scheuchl & Schwenninger (2015). POLAND:

Bogdanowicz et al. (2004). CZECH REPUBLIQUE: Straka et al. (2007). SLOVAKIA: Straka et al. (2007).

SWITZERLAND: Amiet et al. (2004). LIECHTENSTEIN: Bieri (2002). AUSTRIA: Gusenleitner et al. (2012).

SLOVENIA: Gogala (1999). HUNGARY: Józan (2011). NORWAY: Aus t-Ag der, Te l emar k , Vestfo ld, Os t fold,

Oslo (F. Ødegaard, http://artsdatabanken.no/Pages/148154). SWEDEN: Blekinge, Öland, Gotland, Bohuslän

(Janzon et al. 1991). UKRAINE: Romasenko (1995). RUSSIA: Chishmy near Ufa (Tkalců 1974a); southern

European Russia (Osychnyuk et al. 1978).

New records. SPAIN: Girona: 30 km NW Ripoll, 1750 m, 22.7.2011 (leg. J. Halada); Lleida: Cataluña,

Balaguer, Canal d'Urgell, 41°47'01"N 0°49'48"E, 6.8.2009 (leg. J. Smit). ANDORRA: La Massana, 20.7.1999 (leg.

J. Smit). ITALY: Sardinia: Muravera, 2.7.2000 (leg. J. Halada). BULGARIA: Slantschev Brjag, 7.1966 (leg. Z.

Padr); Primorsko env., 6.8.1988 (leg. P. Tyrner); 30 km W Sofia, 12.8.1993 (leg. M. Halada); Trakia, Plovdiv,

20.7.1996 (leg. A. Zaykov); Rodopi, Galabovo, 1.8.1997 (leg. Z. Pedr); Stara Zagora, 5.7.2000 (leg. M. Snizek).

MOLDOVA: Lozova, 15.7.2009 (leg. A. Lozan). TURKEY: Eskisehir: Inönü, 800 m, 1.8.1991 (leg. K. Warncke);

Konya: Güneysinir, Güragaç, 28.7.2000 (leg. M. Kesdek); Nevsehir: Göreme, 1100 m, 25.8.1991 (leg. K.

Warnck e); Erzurum: Agziacik, 20.7.2003 (leg. J.G. Rozen, H. Özbek). RUSSIA: Altai: Chemal, 21.7.2007 (leg. S.

Belokobylskij); Tigirek, 11.7.2012 (leg. M. Shcherbakov); Khakassia: Shira lake, 28.6.2011 (leg. K. Tomkovich).

KAZAKHSTAN: Alma Ata, Medeo, 27.6.1995 (leg. J. Halada); Dshungarskij-Alatau, Rudnitschnij, 44°40'20''N

78°55'38''E, 1200m, 25.7.2002 (leg. M. Kuhlmann). KYRGYZSTAN: Afleatum env., 41.6°N / 71.6°E, 1–3.6.1995

(leg. M. Mücka); southern shore of Issy-Kul lake, Teplokljutschinka, 1650 m, 19.6.1995 (leg. W. Dolin); Ala

Archa, Uzum-Bulat, 5.2000 (leg. V. Gurko); Ala Archa, Kashka-Suu, 1650 m, 7.2000 (leg. V. Gurko); Alai

mountain range, Katla, Karakol, 7.2000 (leg. V. Gurko); Tchatkal mountain range, Khodza-Ata, 41°50'N 71°56'E,

5.7.2000 (leg. Makogonova): Ferghan mountain range, Alash-Too mountains, Alash forest, 8.2000 (leg. V. Gurko);

Osh, Gultcha Ravine, 50 km SSW Gultcha, 39°52'17''N 73°21'26''E, 2530 m, 7.7.2000 (leg. M. Engel).

Distribution. From southern Europe (northern Spain, Andorra, southern France, Italy including Sardinia and

Sicilia) over southeastern Europe (Croatia, Macedonia, Bulgaria, Romania, Moldova) to eastern Turkey and the

Caucasus (Armenia); from western Europe (southernmost United Kingdom, France, Belgium, Netherlands) over

central Europe (Luxembourg, Germany, Poland, Czech Republique, Slovakia, Switzerland, Liechtenstein, Austria,

Slovenia, Hungary), northern Europe (southernmost Norway, southernmost Sweden) and eastern Europe (Ukraine,

Russia) to central Asia (Kazakhstan, Kyrgyzstan) and the Russian Khakassia republic of eastern Siberia. The

species seems to be absent from most of the Iberian Peninsula as well as from Greece. There is a single male from

Crete (Pass near Pinakino, 19.4.1986, leg. W. Vöth, Oberösterreichisches Landesmuseum Linz), which might have

been mislabelled as no other specimens have ever been recorded from this well explored Greek island. The

westernmost record is from Pembrokeshire in southwestern Wales, the northernmost from Oslo province in

southern Norway, the southernmost from Güragaç in southern Konya province of Turkey and the easternmost from

Shira lake in the Khakassia republic of Russia.

Pollen hosts. Oligolectic on Asteraceae (Westrich 1989; Müller 1994). Among the Asteraceae, species of the

Asteroideae (e.g. Anthemis, Aster, Buphthalmum, Inula, Senecio), Carduoideae (e.g. Carduus, Centaurea, Cirsium)

and Cichorioideae (e.g. Cichorium, Crepis, Hieracium, Leontodon, Picris, Tragopogon) are exploited for pollen.

On the Asteroideae, which is probably the most important pollen host taxon among the Asteraceae subfamilies

(Fig. 1), the females take up pollen from the surface of the capitulum directly into their scopa by rapidly moving

the metasoma up and down (“abdominal drumming” sensu Cane 2017). On Carduoideae und Cichorioideae, they

use their hind legs to direct the pollen-bearing flower structures under the seesawing metasoma. To provision one

brood cell, the entire pollen content of about four flower heads of Buphthalmum salicifolium is needed (Müller et

al. 2006).

Nesting biology. Nesting site: The nests are built in empty snail shells of small to medium size (Fig. 5, 6) e.g.

Cepaea, Cernuella, Fruticicola, Helicella, Pomatias, Xerolenta, Zebrina and occasionally also young shells of

Helix (Müller 1994). Nest architecture: The nests contain 1–3, mostly 2 brood cells, which are separated from each

other by partitions consisting of leaf pulp, e.g. from Potentilla or Sanguisorba (Müller 1994). There is no basal

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OSMIINE BEES OF OSMIA (HOPLOSMIA)

wall that seals the innermost brood cell against the rear end of the nest. The shells are sealed at their opening with

an additional wall of leaf pulp. There is an empty vestibule of varying length between nest plug and outermost cell

partition. The latter probably acts as main barrier against predators or parasites as it is distinctly more robust than

the other cell partitions and the nest plug, needing more than 30 flights with leaf pulp for its construction. Nesting

cycle: On average, about 30 foraging bouts are needed to provision one brood cell, which takes about 12.5 h under

good weather conditions (Müller 1994). After having sealed the shell, the female crawls upside down under her

nest and turns it with her legs so that the shell opening is directed tightly towards the ground (Fig. 6), which might

possibly provide some protection against inclement weather. Under good conditions, a female constructs up to 20

brood cells during her flight period, which lasts maximally 10–11 weeks. Interestingly, the females regularly

control their nests up to four weeks after they have sealed them to repair holes and cracks in the nest plug with

newly collected leaf pulp. Osmia spinulosa overwinters as a prepupa in a self-spun cocoon within the brood cell. It

has one generation per year. However, it seems to be a parsivoltine species since about half of all larvae that

emerged from the brood cells were found to undergo metamorphosis only after the second hibernation. Brood

parasites: The megachilid bee species Stelis odontopyga develops as cleptoparasite in the nests of O. spinulosa

(Noskiewicz 1925; Blüthgen 1926). Additional confirmed brood parasites are Chrysura cuprea (Rossi) and C.

trimaculata (Förster) (Chrysididae), Anthrax aethiops (Fabricius) (Bombyliidae), Melittobia acasta (Walker)

(Eulophidae), Pteromalus apum (Retzius) and P. venustus Statz (Pteromalidae) (Kunz 1994; Müller 1994; BWARS

2013).

Behaviour. Male mating behaviour: The males occupy small home ranges, to which they adhere during their

entire flight period, which lasts maximally 5–6 weeks (Müller 1994). Within these home ranges, they search for

females by patrolling Asteraceae flower heads along more or less fixed circular flight routes in a rapid flight, which

is interrupted by short resting periods on the ground. Home ranges and flight routes are never defended against

conspecifics but instead often widely overlap. Sleeping places: The males sleep singly or in small groups within

empty snail shells as do females that have not yet started their nesting activities (Müller 1994). Nesting females

pass the night or periods of bad weather within their nests.

Osmia (Hoplosmia) tyrneri (Tkalců 1992)

Hoplosmia (Hoplosmia) tyrneri Tkalců 1992: 220. Type material: Holotype ♂, “Tashkent, Galvasai, 25.6.1980” (Uzbekistan),

Oberösterreichisches Landesmuseum Linz.

Literature records. UZBEKISTAN: Chatkal Mountains, 70 km NE Tashkent/Galvasai (Tkalců 1992).

Distribution. Known only from the sourroundings of Tashkent in eastern Uzbekistan.

Nesting biology. Unknown.

Pollen hosts. Unknown.

Notes. Female unknown. B. Tkalců erroneously labelled the specimen that was designated as holotype in his

original publication as a paratype, which is deposited in the Oberösterreichische Landesmuseum Linz. Thus, Linz

is actually the holotype depository of O. tyrneri and not the Tyrner Collection in Litvinov as stated in Tkalců

(1992). The males of O. tyrneri and O. spinulosa differ only slightly, e.g. in the shape of the sternal spine, the

punctation of the integument and the colour of the body pilosity (see Key to species), suggesting conspecificity.

However, these separating characters do not seem to clinally change from west to east, since males of O. spinulosa

from both northern Kyrgyzstan and southeastern Kazakhstan are morphologically almost identical to those from

Europe. Therefore, O. tyrneri is regarded here as a species of its own in spite of its very close resemblance to O.

spinulosa.

Osmia bidentata species group

The members of this species group are characterized by the shape and punctation of the female clypeus and the nest

building material. In contrast to the other O. (Hoplosmia) species, the clypeus lacks an impunctate apical margin, it

is apically bent rather than flattened and its punctation is apically composed of both small and large punctures

rather than of uniformly fine punctures. Brood cell partitions and nest plug are constructed from mud, whereas all

the other O. (Hoplosmia) species use leaf pulp as nest building material.

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312

Osmia (Hoplosmia) anceyi Pérez 1879

Osmia Anceyi Pérez 1879: 187. Type material: Lectotype ♀, by designation of Tkalců (1974a), “Environs de Marseille”

(France), Muséum National d’Histoire Naturelle Paris.

Literature records. FRANCE: Alpes-de-Haute-Provence, Bouches-du-Rhône, Pyrénées-Orientales (Benoist

1931); Bouches-du-Rhône, Hérault, Pyrénées-Orientales (Benoist 1931, as Osmia bidentata). SPAIN: Alicante,

Sevilla, Granada, Barcelona, Burgos, Huesca, Ciudad Real (Ceballos 1956, as Osmia bidentata); Escorial/Alt

Castilion/Venta de Baños (Tkalců 1974a); Cataluña/Zariquiey (Tkalců 1974a). PORTUGAL: Elvas (Tkalců

1974a). TUNISIA: Tunis (Alfken 1928).

New records. SWITZERLAND: Valais: Zeneggen, 28.6.1987, 13.6.1993 (leg. A. Müller); Hohtenn, 29.6.1990

(leg. A. Müller); Baltschieder, Lengmüra, 700–800 m, 27.7.1997 (leg. A. Müller); Visperterminen, 14.8.1997 (leg.

W. Arens); Erschmatt, 1300–1500 m, 8.6.2003 (leg. A. Müller). FRANCE: Drôme: Bollène, Rochegude, 2.7.2005

(leg. J. Smit); Alpes-de-Haute-Provence: Riez, 3.9.1977 (leg. G. Le Goff); Gréoux-les-Bains, 18.6.1978 (leg. G. Le

Goff); 30 km S Digne-les-Bains, 570 m, 18.7.2011 (leg. J. Halada); Vaucluse: Avignon, 2.8.1957 (leg. W. Schläfle);

Robion, 28.8.1974 (leg. G. Le Goff); Saumane, 1.7.1997 (leg. G. Le Goff); Bédoin, 13 km NE Carpentras,

18.7.2002 (leg. J. van der Smissen); Var: W Grasse, Lac de St. Cassien, 1.7.1999 (leg. S. Risch); Bouches-du-

Rhône: Salon-de-Provence, Lamanon, 20.7.2000 (leg. J. Smit); Pyrénées-Orientales: Le Barcarès, 13.7.1992 (leg.

G. Le Goff); Saillagouse, 1345 m, 22.7.2001 (leg. M. Vandenbergh); 10 km S Prades, 20.7.2011 (leg. J. Halada);

Corsica: Verghia, 2.6.2001 (leg. G. Le Goff). SPAIN: Girona: Puigcerdà, 14.7.1963 (leg. H. Hamann); Tarragona:

Ginestar, 50 km N Tortosa, 4.5.2000 (leg. E. Scheuchl); Huesca: Loarre, 12.6.1994 (leg. F. Amiet); Burgos: Lerma,

rio Arlanza, 5.7.2006, 9.7.2007 (leg. J. Smit); Segovia: Hontanares Eresma, Sierra de Guadarrama, 920 m,

29.7.2008 (leg. F.J. Ortiz-Sanchez); Alicante: Guardamar del Segura, 16.5.2001 (leg. G. Le Goff); Huelva:

Galaroza, Jabugo, 650 m, 17.6.2015 (leg. J. Smit); Almeria: La Aldeilla, El Ejido, 60 m, 27.5.2005 (leg. F.J. Ortiz-

Sanchez); Berja, Sierra de Gádor, 500 m, 10.5.2006 (leg. F.J. Ortiz-Sanchez). MOROCCO: Fès-Meknès: Ifkern, 25

km E Boulemane, 25.5.1995 (leg. M. Halada); Achlouj, Jnane, 16.6.2013 (leg. L. Castro, A. Francois);

Casablanca-Settat: between Boulaouane and Tizi Ouchen, 900 m, 20.5.1998 (leg. P. Rasmont); Marrakech-Safi: El

Garma, Kebdana-Marrakech, 5.1972 (leg. A. Pardo); Marrakech, Takatert, 870 m, 6.7.1996 (leg. P. Rasmont);

Marrakech, El Caid Omar, 600 m, 7.6.1996 (leg. P. Rasmont); Drâa-Tafilalet: SE Tazenakht, 12.5.2003 (leg. M.

Snizek); Souss-Massa: 30 km SWW Tata, Imitek, 13.4.2015 (leg. C. Schmid-Egger). ALGERIA: Constantine:

Cons, Ben badis, 15.5–4.6.2008 (leg. S. Aguib); Mila: Ouledbazer, 28.4–26.5.2013 (leg. S. Aguib); Guelma:

Guelma, Campus, 17.5–10.6.2008 (leg. S. Aguib). TUNISIA: Tuni s: Tunis, 4.1930 (leg. R.Meyer).

Distribution. Switzerland (Valais), southern France (including Corsica), Iberian Peninsula (Spain, Portugal),

Maghreb (Morocco, Algeria, Tunisia). Osmia anceyi seems to be entirely absent from Italy, even in the country’s

westernmost parts (G. Pagliano personal communication).

Subspecific classification. The apical margin of the clypeus of females occurring in Morocco, Algeria and

Tunisia is slightly less emarginate than in females from southwestern Europe. Based on this difference, Zanden

(1994) established two subspecies, i.e. Osmia anceyi anceyi Pérez 1879 distributed in southwestern Europe and O.

anceyi biarmica (Zanden 1994) ranging from Morocco to Tunisia.

Pollen hosts. Oligolectic on Asteraceae (based on 26 pollen loads from 18 different localities in Switzerland,

France, Spain, Morocco and Algeria and on field observations). All 26 pollen loads exclusively consisted of pollen

of Carduoideae (Centaurea and/or thistles), suggesting that O. anceyi restricts pollen harvesting to this subfamily.

Flower records: Carduus nutans, Centaurea algeriensis, C. calcitrapa, C. jacea, C. paniculata, C. solstitialis, C.

sphaerocephala, Onopordum illyricum (label records).

Nesting biology. The nests are built in various preexisting cavities: insect burrows in dead wood, dead hollow

plant stems, burrows in the pith of dead stems (e.g. Asphodelus) excavated by other aculeates (e.g. Ceratina) or

abandoned nests of aculeate hymenopterans in clay walls (Alfken 1928; Torres et al. 1997; Amiet et al. 2004; G. Le

Goff personal communication; A. Müller unpublished data). The nests contain up to nine linearly arranged brood

cells, which are separated from each other by partitions constructed from mud, which is sometimes mixed with

small particles of pith. Coelioxys osmiae Alfken is assumed to be a brood parasite of O. anceyi (Alfken 1928).

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OSMIINE BEES OF OSMIA (HOPLOSMIA)

Osmia (Hoplosmia) bidentata Morawitz 1876

Osmia bidentata Morawitz 1876: 38. Type material: Syntypes ♂♂, ♀♀, “Etschmiadzin” (Armenia), type depository unknown.

Osmia affinis Frivaldsky 1877: 360. Type material: Syntypes ♂♂, ♀♀, “In Comitatu Crassoviensi ad Oraviczam, in Comitatu

Pestiensi vero ad Budapestinum et in Praedio-Peszér” (Hungary), type depository unknown. Synonymy by Mocsáry

(1884).

Literature records. AUSTRIA: Styria, Lower Austria, Vienna, Burgenland (Gusenleitner et al. 2012); Vienna,

Nordbahnhof (Zettel et al. 2016). CZECH REPUBLIQUE: Moravia: Znojmo, Kobyli, Pouzdřany, Dolní

Věstonice, Pavlovské vrchy, Uherské Hradiště (Tkalců 1974a). SLOVAKIA: Trenčín, Štúrovo, Gbelce, Mužla,

Kamenica n. Hr., Chl’aba, Turňa n. Bodvou, Košice (Tkalců 1974a). HUNGARY: Great Hungarian Plain,

Transdanubian Hills, Transdanubian Mountains, North Hungarian Mountains, West Hungary (Józan 2011).

POLAND: Southeastern Poland (Bogdanowicz et al. 2004). ROMANIA: Agigea, Runc, Racova, Urechesti,

Comorova, Ulea, Cordun, Traian, Piatra Neamt, Gugesti (Ban-Calefariu 2009, Tomozii & Toma 2011). UKRAINE:

Crimea, Zaporozhzhie, Donetzk, Luhansk (Romasenko 1995). RUSSIA: southern European Russia (Osychnyuk et

al. 1978). ITALY: Piemonte, Lombardia, Friuli-Venezia Giulia, Veneto, Lazio, Abruzzi (Comba & Comba 1991,

Pagliano 1994). SLOVENIA: Prealpine region (Gogala 2014). CROATIA: Brač Island/S. Pietro-Neresi (Maidl

1922); Zadar-Sukosany, Split (Tkalců 1974a); Briševo, Gorica, Kakma, Karlobag, Lukovo Šugarje, Rijeka, Tribanj

Krušćica (Józan 2009). SERBIA: Novi Sad (Tkalců 1974). MONTENEGRO: Ulcin (Tkalců 1974a).

MACEDONIA: Dojran lake (Tkalců 1974a); Tetovo (Zanden 1984a). ALBANIA: Pashtrik (Maidl 1922).

BULGARIA: Sandanski, Slantschev Brjag, Aitos (Tkalců 1974a). TURKEY: Izmir, Balikesir, Istanbul, Aydin,

Antalya, Nevsehir, Karaman, Sivas, Bayburt, Erzurum, Bitlis, Ardahan, Kars, Agri, Van (Tkalců 1979, Özbek &

Zanden 1992); Ankara, Cankiri, Eskisehir, Yozgat, Sivas (Güler & Cagatay 2006). IRAN: Teheran province/road

Sar Ziarat-Chalus (Nadimi et al. 2013). SYRIA: Barze, Damas (Zanden 1989). ISRAEL AND PALESTINE:

Jerusalem (Tkalců 1974). EGYPT: Cairo (Tkalců 1974a, 1979).

New records. AUSTRIA: Lower Austria: Mistelbach, 17.7.1995 (leg. K. Dollfuss); Burgenland: Winden am

See, 14.8.1984 (leg. M. Schwarz). HUNGARY: Buda, 7.7.1886 (leg. H. Friese). ITALY: Tos c a na : Massa

Marittima, 600 m, 27.7.2005 (leg. A. Müller); Abruzzo: Massa d’Alba, 900 m, 23.8.2002 (leg. A. Müller); Puglia:

Mte. Gargano, Monte Sant’Angelo, 530 m, 30.6.1994 (leg. A. Müller). CROATIA: Makarska, 27.6.2011 (leg. Z.

Pedr). SERBIA: Pristina, 20.6.2000 (leg. Ø. Berg). MACEDONIA: Ohrid, 24.7.1958 (leg. W. Schläfle). GREECE:

Western Macedonia: Kozani, Proastio, 620 m, 18.6.1990 (leg. J. Tiefenthaler); Central Macedonia: Saloniki,

6.1960 (leg. W. Schläfle); Central Greece: Orchomenos, Arkadia, 25.6.1996 (leg. W. Arens); Western Greece:

Patras, Panahaikon mountains, 800–1700m, 25.6.1998 (leg. W. Arens); Peloponnese: Epidauros, Limera, 9.6.1996

(leg. W. Arens); Crete: Kato Horio NE Ierapetra, 15.4.1986 (leg. W. Vöth); Aegean Islands: Lesbos, Polichitos,

24.6.2008 (leg. C. Sedivy); Thasos, Astris, 21.6.2012 (leg. M. de Courcy); Syros, Komito, 11.6.2013 (leg. I.

Vavitsas); Kea, Poisses-Koundouros, 27.6.2013 (leg. T. Petanidou). BULGARIA: Kresna-Piria, 5.1967 (leg. M.

Kocourek); Arkutino, 7.1968 (leg. A. Görtier); Slantschev Brjag, 6.1972 (leg. M. Kocourek); Neseban, 7.1986 (leg.

Z. Pedr); Szozopol, 8.7.1986 (leg. Kolacek); Primorsko env., 6.8.1988 (leg. P. Tyrner); SW Razlog, 16.8.1993 (leg.

Z. Pedr); Sandanski, 6.1994 (leg. M. Kocourek); Trakia, Plovdiv, 20.7.1996 (leg. A. Zaykov); Rodopi, Studenec,

25.7.1997 (leg. Z. Pedr); Dervinska Mogila, 25 km NE Svilengrad, 500m, 20.6.2008 (leg. M. Halada). TURKEY:

Canakkale: 6 km N Ezine, 35 m, 27.7.2006 (leg. M. Halada); Izmir: 10 km NE Odemis, 1200 m, 3.7.2006 (leg. M.

Halada); Aydin: Adnan Menderes University, Campus, 4.7.2006 (leg. E. Scheuchl); Manisa: 15 km SEE Salihli,

170 m, 2.7.2006 (leg. M. Halada); Denizli: around Denizli, 14.6.2006 (leg. E. Scheuchl); Eskisehir: Porsuk Baraji

Sebran, 8.7.1993 (leg. K. Denes); Burdur: 20 km SW Burdur, 940 m, 7.7.2006 (leg. J. Halada); Isparta: Egirdir

Gölu, 5 km N Akkecili, 920 m, 10.7.2006 (leg. M. Halada); Konya: Çumra, 1020 m, 12.8.2000 (leg. M. Kesdek);

Aksaray: Esmekaya, 52 km W Aksaray, 16.7.1998 (leg. C. Schmid-Egger); Nevsehir: Göreme, 23.6.1993 (leg. M.

Halada); Nigde: Camardi, 13.7.1997 (leg. M. Halada); Kayseri: Burhaniye, 1320 m, 13.7.1996 (leg. P. Rasmont);

Sivas: Karagöl, 1260 m, 14.7.1996 (leg. P. Rasmont); Malatya: Erkenek, 80 km SW Malatya, 9.7.1997 (leg. M.

Halada); Adiyaman: Nemrut Dagi Karadut, 2.7.1993 (leg. K. Denes); Erzincan: Erzincan, 18.7.1997 (leg. E.

Yildirim); Elazig: Hazar Gölü SE Elazig, 29.6.2000 (leg. M. Halada); Sanliurfa: Caylarbasi, 70 km N Urfa,

2.6.1998 (leg. M. Halada); Erzurum: Eskipalat, 1810 m, 20.7.1997 (leg. P. Rasmont); Bingöl: Solhan-Baglan, 2120

m, 25.6.2004 (leg. M. Kesdek); Mardin: 20 km N Mardin, 21.6.1997 (leg. M. Halada); Bitlis: 20 km SW Bitlis,

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23.6.1997 (leg. M. Halada); Igdir: 20 km NW Igdir, 29.6.1997 (leg. M. Halada); Agri: Gökoglu, 1680 m, 22.7.1997

(leg. P. Rasmont); Va n : Tevekli, 1740 m, 23.7.1997 (leg. P. Rasmont). ARMENIA: Armawir: Vanand, 1120 m,

18.6.2006 (leg. Ø. Berg). IRAN: East Azerbaijan: Sis, 10 km E Shabestar, 1540 m, 19.6.2012 (leg. Z. Pedr);

Kohgiluyeh and Boyer-Ahmad Province: Kuh Go near Sisakht, 2500 m, 9.6.2010 (leg. M. Halada); Fars: Komehr,

2380 m, 9.6.2010 (leg. M. Halada); Lorestan: 10 km SW Dorud, 1520 m, 27.5.2014 (leg. J. Halada). SYRIA:

Khabab, 60 km S Damascus, 14.5.1996 (leg. M. Halada); 30 km S Suwayda, 15–17.5.1996 (leg. M. Halada); Dara

Nawa, 18.5.1996 (leg. M. Halada); 40 km NE Damascus, 22.5.1996 (leg. M. Halada); Dibbin, 30 km W Damascus,

19.6.2000 (leg. M. Halada); Kafr, Suwayda, 21.6.2000 (leg. M. Halada). ISRAEL AND PALESTINE: Northern

District: Golan, 1 km S Ein Kinya, 600 m, 5.5.2010 (leg. C. Sedivy, C. Praz); Mt. Hermon, 2180 m, 27.6.2010 (leg.

A. Dorchin); Upper Galilee, Nahal Ga'ton 1.4 km N Ga'ton, 240 m, 30.5.2011 (leg. A. Dorchin); Upper Galilee,

Har Meron, 14.6.2013 (leg. G. Pisanty); West Bank: Wadi Og, -10 m, 7.4.2012 (leg. J.S.Ascher, A.Payne); Nahal

Perat, 14.3.2015 (leg. G. Pisanty). JORDAN: Jordan Valley, Tall al Arbatin, 20 km S North Shuna, 19.4.1996 (leg.

M. Halada); Jordan Valley, South Shuna, 25–26.4.1996 (leg. M. Halada); Jordan Valley, Dayr Alla, 27.4.1996 (leg.

M. Halada); 30 km N Tafila, 2.5.1996 (leg. M. Halada); N Zarga, 26.4.2002 (leg. M. Snizek); 15 km S of Madaba,

28.4.2006 (leg. F. Kantner); Dead Sea, road Al-Tafila-Feifa, 18.4.2007 (leg. C. Praz, C. Sedivy, A. Müller); Wadi

Mujib, King’s Highway, 19.4.2007 (leg. C. Praz, C. Sedivy, A. Müller).

Distribution. From eastern Austria, southern Czech Republique, Slovakia and Hungary over southeastern

Poland and Romania to southern Ukraine and southern European Russia; from Italy and Slovenia over the Balkan

peninsula (Croatia, Serbia, Montenegro, Macedonia, Albania, Greece including Crete, Bulgaria) and Turkey to the

Caucasus (Armenia) and central Iran; from Turkey over the Levant (Syria, Israel and Palestine, Jordan) to northern

Egypt. The westernmost record is from Mondovi in Italian Piemonte province (G. Pagliano personal

communication), the northernmost from Uherské Hradiště in southern Moravia, the southernmost from Cairo in

northern Egypt and the easternmost from Komehr in Iranian Fars province. The species’ occurrence in

Turkmenistan, Mongolia and the Russian Far East as reported by Romankova (1995) appears to be doubtful and

needs verification. Osmia bidentata seems to be entirely absent from France, the Iberian Peninsula and the

Maghreb, where it is replaced by O. anceyi as already assumed by Tkalců (1979).

Subspecific classification. The metasomal scopa of females occurring on Lesbos, in Turkey, Iran, the Levant

and Egypt is whitish rather than yellowish-red as in females of the more western populations and of the Caucasus.

Based on this difference, Tkalců (1979) established two subspecies, i.e. Osmia bidentata bidentata Morawitz 1876

distributed mainly in Europe and O. bidentata pallens (Tkalců 1979) ranging from easternmost Europe over

Turkey eastwards to Iran and southwards to northern Egypt.

Pollen hosts. Oligolectic on Asteraceae (based on 62 pollen loads from 27 different localities in Italy, Austria,

Hungary, Macedonia, Greece, Turkey, Syria, Israel and Jordan and on field observations). All loads exclusively

consisted of Centaurea pollen except for two loads that additionally contained moderate quantities of thistle pollen,

indicating that O. bidentata most probably restricts pollen harvesting to the subfamily Carduoideae (Fig. 3). One

load contained small amounts of Onobrychis pollen (Fabaceae) beside Centaurea pollen, suggesting that the

females very rarely also exploit pollen hosts other than the Asteraceae. These findings are in contrast to Güler &

Sorkun (2007), who assume O. bidentata to be polylectic, harvesting pollen from flowers of eleven plant families.

Flower records: Carduus spec., Centaurea stoebe (Tkalců 1974a); Arctium, Carduus spec., Centaurea iberica, C.

spinosa, Cirsium spec., Onopordum spec. (label records).

Nesting biology. The nests are built in various preexisting cavities, such as insect burrows in dead wood, dead

hollow plant stems (e.g. Phragmites) or cavities in the soil (Popovici-Baznosanu 1909; Banaszak & Romasenko

2001; A. Müller unpublished data), possibly also in abandoned nests of aculeates in loess scarps (H. Wiesbauer

personal communication). They contain 1–11 linearly arranged brood cells, which are separated from each other by

partitions made of mud. In Phragmites stems, the entire brood cells including their walls may be constructed from

mud. The nest is sealed by a one-layered wall of mud at its opening. The space between the outermost cell and the

nest plug is 1.5–5 cm long and usually empty, but is occasionally divided up by an additional wall of mud. Osmia

bidentata overwinters in the prepupal stage (Popovici-Baznosanu 1909).

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OSMIINE BEES OF OSMIA (HOPLOSMIA)

Osmia ligurica species group

This species group unites all O. (Hoplosmia) species, which i) have the metanotum not overhung by the scutellum,

ii) are characterized by an apically flattened and uniformly punctured clypeus, iii) nest in preexisting cavities other

than snail shells and iv) use leaf pulp as nest building material.

Osmia (Hoplosmia) dido Gribodo 1894

Osmia dido Gribodo 1894: 289. Type material: Holotype ♀, “Algeria” (Algeria), Museo Civico di Storia Naturale Genova.

Osmia abbreviata Pérez 1895: 13. Nomen praeoccupatum (not Osmia abbreviata Morawitz 1875). Type material: Lectotype ♀,

by designation of Tkalců (1974a), “Algéria” (Algeria), Muséum National d’Histoire Naturelle Paris.

Osmia compacta Pérez 1896: not paginated single page appendix to Pérez (1895). Nomen novum with same type specimen for

preoccupied Osmia abbreviata Pérez 1895 (not Osmia abbreviata Morawitz 1875). Synonymy in Zanden (1990).

Literature records. ALGERIA: Algiers (Alfken 1914); Oran/Ain Zaatout/Ouréz mountains, Bouira, Tipasa

(Tkalců 1974a). TUNISIA: Tunis (Tkalců 1974a).

New records. MOROCCO: Fès-Meknès: 15 km SE Sefrou, 26–27.5.1995 (leg. M. Halada); Bhalil, 10 km NW

Sefrou, 28.5.1995 (leg. M. Halada); SW of Sefrou, 16.5.2003 (leg. M. Halada). ALGERIA: Biskra: El Kantara,

17.5.2008 (leg. S. Aguib). TUNISIA: Bizerta: Ghar El Melh E Bizerta, 26.5.1999 (leg. O. Niehuis); Zaghouan: 30

km SE Zaghouan, 19.4.1981 (leg. M. Schwarz); Kef: 5 km N El Kef, 22.6.1994 (leg. M. Hauser); Sfax: 30 km SW

Sfax, 18.4.1981 (leg. M. Schwarz).

Distribution. Maghreb (Morocco, Algeria, Tunisia).

Pollen hosts. Probably oligolectic on Asteraceae (based on eight pollen samples from two different localities

in Morocco). As all pollen loads analysed were either pure loads of Centaurea (n = 6) or contained pollen of both

Centaurea and thistles (n = 2), O. dido is likely a specialist of Carduoideae. Flower record: Centaurea nicaeensis

(label record).

Nesting biology. Unknown.

Osmia (Hoplosmia) distinguenda (Tkalců 1974)

Anthocopa (Odontanthocopa) distinguenda Tkalců 1974a: 129. Type material: Holotype ♂, “Jerusalem, 31.5.1929” (Israel),

Natural History Museum London.

Literature records. TURKEY: Konya, Mersin, Kayseri, Sivas, Malatya, Mardin, Erzurum, Mus, Hakkari, (Özbek

& Zanden 1992).

New records. GREECE: Western Greece: Patras, Panachaiko mountains, 800–1700 m, 25.6.1998 (leg. W.

Arens); Peloponnese: Cape Tenaro, Mani, 20 m, 16.5.1995, 4.5.2000 (leg. W. Arens), 23.4.2001 (leg. A. Müller);

TURKEY: Denizli: 35 km SSE Denizli, 970 m, 5.7.2006 (J. Halada); Kütahya: 20 km NEE Kütahya, 13.7.2006

(leg. J. Halada); Isparta: Karakus Dagi centr., 1460 m, 11.7.2006 (leg. J. Halada); Eskisehir: Porsuk Baraji Sebran,

8.7.1993 (leg. M. Halada); Mersin: Kiskalesi near Silifke, 9–13.5.1988 (leg. N. Mohr); Sivas: 20 km E Gurun,

Mezikiran Gecidi, 10.7.1997 (leg. M. Halada); Kahramanmaras: Kahramanmaras env., 601 m, 18.5.2006 (leg.

Hazir, Cobanoglu); Gaziantep: 10 km W Gaziantep, 20.6.1997 (leg. M. Halada); Adiyaman: Nemrut Dagi,

Karadut, 2.7.1993 (leg. M. Halada); Sanliurfa: Birecik S Halfeti, 30.5.1998 (leg. M. Snizek); Mardin: 20 km N

Mardin, 21.6.1997 (leg. M. Halada); Erzurum: Umudum Yavl., 2800 m, 8.8.1991 (leg. H. Özbek); Bitlis: Kokabsu,

27.7.1978 (leg. H. Özbek); Kars: Pasli, 50 km S Kars, 1.7.1997 (leg. M. Halada); Va n : 10 km N Muradiye,

27.6.1993 (leg. M. Halada). ARMENIA: Armavir: Vanand, 1120 m, 18.6.2006 (leg. Ø. Berg). SYRIA: Khabab, 60

km S Damascus, 14.5.1996 (leg. M. Halada); Dibbin, 30 km S Suwayda, 15–17.5.1996 (leg. M. Halada); 30 km N

Dara Nawa, 18.5.1996 (leg. M. Halada); Anata, 50 km S Suwayda, 20–21.5.1996 (leg. M. Halada); 40 km NE

Damascus, 22.5.1996 (leg. M. Halada); 50 km S Homs, 24.5.1996 (leg. M. Halada); Tartus, Oal at al-Hisn,

8.6.2000 (leg. K. Denes); 30 km W Damascus, 19.6.2000 (leg. M. Halada). ISRAEL AND PALESTINE: Northern

District: Golan, 1 km S Ein Kinya, 5.5.2010 (leg. C. Sedivy, C. Praz); Mt. Hermon, 2180 m, 28.06.2010 (leg. A.

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Dorchin); Lake Tiberias, 2 km NNE Tiberias, 70 m, 25.5.2011 (leg. S. Risch); Lower Galilee, W Moreshet, 250 m,

28.5.2011 (leg. S. Risch); Upper Galilee, 1.8 km SE Bet Rimon, 220 m, 28.5.2011 (leg. A. Dorchin, S. Risch); 1

km SW Ziv'on, 15.5.2015 (leg. G. Pisanty); Nabi Hazuri, 26.6.2015 (leg. G. Pisanty); Haifa District: 40 km NE

Haifa, 1 km E Hurfeish, 15.5.1996 (leg. C. Schmid-Egger); Mt. Carmel, Hai Bar, 29.5.2000, 12.6.2000 (leg. M.

Török); Jerusalem District: Judean Foothills, Har'el, 22.5.2009 (leg. G. Pisanty); Judean Foothills, Beit Govrin,

28.3.2010 (leg. G. Pisanty); Judean Foothills, Ya'ar Yish'I, 21.4.2010 (leg. T. Koznichki); Judean Mountains, Nataf,

16.5.2012 (leg. Y. Berner); Judean Foothills, Park Britannia, 12.4.2013 (leg. Y. Berner), 7.5.2015 (leg. T.

Chaprazaro); 1 km N Nahshon, 30.5.2015 (leg. G. Pisanty); Westbank: Judean Desert, Za’tara, 5.5.2014 (leg. I.

Arar). Southern District: Judean Foothills, Lakhish, 6.5.2012 (leg. Y. Berner); JORDAN: Jordan Valley, South

Shuna, 25–26.4.1996 (leg. M. Halada); Jordan Valley, Mubalath, 27.4.1996 (leg. M. Halada); 30 km N Tafila,

2.5.1996 (leg. M. Halada); 10 km N Petra, 3.5.1996 (leg. M. Halada); Jordan Valley, 20 km S North Shuna, Tall Al

Arbatin, 19.6.1996 (leg. M. Halada); Kafr, Suwayda, 21.6.2000 (leg. M. Halada); Al Karak env., 16.4.2002 (leg.

M. Snizek); 10 km N Jerash, 20.4.2002 (leg. M. Snizek); 30 km NW Ajlun, 600m, 29.4.2006 (leg. K. Denes); NW

Pella env., -80 m, 29.4.2006 (leg. K. Denes).

Distribution. From the Peloponnese peninsula eastwards to eastern Turkey and Armenia and from Turkey

southwards to the Levant (Syria, Israel and Palestine, Jordan).

Pollen hosts. Oligolectic on Asteraceae (based on 34 pollen samples from 19 different localities in Greece,

Turkey, Syria, Israel and Jordan and on field observations). As all pollen loads analysed were either pure loads of

Centaurea (n = 23) and thistle pollen (n = 5) or contained pollen of both Centaurea and thistles (n = 6), O.

distinguenda probably restricts pollen harvesting to the Carduoideae. Flower records: Centaurea crocodylium, C.

hyalolepis, C. iberica (label records).

Nesting biology. Observations from both Israel and southern Greece indicate that the brood cells are built

singly or in small groups within small cavities of rocks and stones (G. Pisanty and H. Wiesbauer personal

communication; A. Müller unpublished data; Fig. 8). They consist of leaf pulp with the rock surface usually

forming part of the cell walls. The nest cavity is closed by a wall made of leaf pulp.

Osmia (Hoplosmia) hermonensis (Tkalců 1992)

Hoplosmia (Odontanthocopa) hermonensis Tkalců 1992: 223. Type material: Holotype ♂, “Mt. Hermon, 1700 m, 11.6.1976”

(Israel), Naturalis Biodiversity Center Leiden.

Distribution. Only known from the type series collected at Mount Hermon in northernmost Israel.

Pollen hosts. Unknown.

Nesting biology. Unknown.

Note: Female unknown.

Osmia (Hoplosmia) larochei Tkalců 1993

Osmia (Odontanthocopa) larochei Tkalců 1993: 810. Type material: Holotype ♀, “Gran Canaria Cueva Grande Las Palmas,

18.6.1984” (Spain: Canary Islands), La Roche Collection.

Diagnosis of the hitherto unknown ♂: The male of O. larochei is morphologically very close to O. olgae and O.

scutellaris. It differs from O. olgae by the distinctly longer pilosity of the mesepisternum, the wider scutellum and

the dark rather than reddish-brown tergum 7 (see Key to species). Characters distinguishing it from O. scutellaris

are the shape of the preapical lobes of tergum 6, the form of the labrum and the shape of sternum 4 (see Key to

species).

New records: SPAIN: Canary Islands: Gran Canaria, Los Pechos, 18.3.1988 (leg. Scaramozzino).

Distribution. Restricted to the Canary Islands and known only from Gran Canaria.

Pollen hosts. Unknown. Flower record: Andryala pinnatifida (Hohmann et al. 1993).

Nesting biology. Unknown.

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OSMIINE BEES OF OSMIA (HOPLOSMIA)

Osmia (Hoplosmia) ligurica Morawitz 1868

Osmia ligurica Morawitz 1868: 150. Type material: Syntypes ♂♂, ♀♀, “Bei Nizza im Thale des Paglione” (France), type

depository unknown.

Osmia detrita Pérez 1879: 188. Type material: Lectotype ♀, by designation of Tkalců (1974a), “Bordeaux, Marseille” (France),

Muséum National d’Histoire Naturelle Paris. Synonymy in Schmiedeknecht (1886).

Literature records. SPAIN: Barcelona (Ceballos 1956); FRANCE: Gironde, Pyrénées-Orientales, Gard, Vaucluse,

Bouches-du-Rhône, Var, Alpes-Maritimes (Benoist 1931); Vaucluse, Bouches-du-Rhône, Var (Tkalců 1974a);

Corsica (Zanden 1980). SWITZERLAND: Geneva (Amiet et al. 2004). AUSTRIA: Lower Austria (Gusenleitner et

al. 2012). SLOVAKIA: Nitra, Štúrovo, Plešivec, Turňa n. Bodvou (Tkalců 1974a, Straka et al. 2007). HUNGARY:

Transdanubian Hills, Transdanubian Mountains, West Hungary (Józan 2011). ROMANIA: Ban-Calefariu (2009).

ITALY: Umbria, Campania (Tkalců 1974a); Piemonte, Liguria, Emilia-Romagna, Toscana, Lazio, Abruzzi, Molise,

Campania, Puglia, Basilicata, Sicily, Sardinia (Pagliano 1994). SLOVENIA: Portorož (Tkalců 1974a);

submediterranean region (Gogala 2014). CROATIA: Funtana, Krk, Rijeka, Sveta Jelena, Vela Učka (Józan 2009).

MONTENEGRO: Ulcin (Tkalců 1974a). MACEDONIA: Ohrid (Zanden 1984a). ALBANIA: Tkalců (1974a).

GREECE: Corfu (Tkalců 1974a). BULGARIA: Slantschev Brjag (Tkalců 1974a). TURKEY: Mersin (Zanden

1980); Erzincan, Konya, Icel (Özbek & Zanden 1992). ARMENIA: Osychnyuk et al. (1978). IRAN, Fars:

Nurabad/Durahi (Khodaparast & Monfared 2013).

New records. MOROCCO: Tanger-Tétouan-Al Hoceïma: Chefchaouen, 730 m, 4.5.2002 (H.J. Flügel); Fès-

Meknès: El-Menzel, 30 km E Sefrou, 29.5.1995 (leg. M. Halada). PORTUGAL: Braga: Fafe, San Jorge , 25.5.1991

(leg. M. Schwarz); Castelo Branco: Serra Estrella, 1450 m, 16.7.2009 (leg. V. Bartak); Lisbon: Sobreda, 5.5–

24.5.1985 (leg. S. Risch); Faro: Carrapateira, 20 km SSW Aljezur, 13.5.1995 (leg. J. Gusenleitner). SPAIN:

Girona: Pirineos Orientales, 30 km NW Ripolli, 1750 m, 22.7.2011 (leg. J. Halada); Barcelona: Parc Natural del

Garraf, 19.5.2010 (leg. J. Bosch); Tarragona: Ginestar, 50 km N Tortosa, 4.5.2000 (leg. E. Scheuchl); Castellon:

Castellon env., 6.1997 (leg. Kadlecova); Valencia: Valencia env., 1.5.1997 (P. Stary); Alicante: Moraira, 15 km

SSW Xabla, 3.5.2001 (leg. E. Scheuchl); Murcia: 25 km SW Cartagena, 12.5.2003 (leg. M. Halada); Jaén: Sierra

Pozo, Mt. Palomas, 1450 m, 11.6.2003 (leg. M. Kafka); Sevilla: W Villamanrique de la Condesa, 10.5.2013 (leg. J.

Smit); Huelva: E El Rocio, Coto Donana, National Park, Arroyo de Partido, 19.4.2003 (leg. S. Roberts); Almeria:

Sierra Filabres, 9 km E Albanchez, 23.4.2003 (leg. M. Halada); Granada: Sierra de Tejeda, Punte de Salina, 700 m,

5.2003 (leg. F. Kantner); Malaga: Parauta 13 km S Ronda, 800 m, 5.10.2004 (leg. S. Risch); Cadiz: Chiclana 23

km S Cadiz, 8.4.2010 (leg. C. Schmid-Egger); Menorca: 2007 (leg. P. Mendez). FRANCE: Drôme: Plateau du

Coiron, Sceautres-Taverner, 16 km NW Montélimar, 12.7.2002 (leg. J. van der Smissen); Vauc l us e : 5 km N

Roussillon, Lioux, 9.6.1997 (leg. O. Niehuis); Alpes Maritimes: La Bollène-Vésuble, 630 m, 12.7.2010 (leg. C.

Schmid-Egger); Var : Le Muy, L'Endre, 50 m, 15.5.2006 (leg. Hendrichx); Gard: Nîmes, Marguerittes, 7.5.2000

(leg. E. Scheuchl); Aude: 10 km S Carcassone, 85 km SW Toulouse, 18.5.2003 (leg. M. Halada). ROMANIA:

Moldova Noua, 27.5.2002 (leg. M. Halada). ITALY: Friuli Venezia Giulia: Gemona di Friule, 230 m, 19.6.2006

(leg. H.J. Flügel); Valle d'Aosta: Parleaz, 1250 m, 2.6.1999 (leg. E. Scheuchl); Emilia Romagna: Berceto,

15.7.1988 (leg. Zinnert); Liguria: Levanto, 31.5–5.6.1999 (leg. M. Herrmann); Toscana: Monte Argenario, 40 km

S Grosseto, 5.5.1995 (leg. C. Schmid-Egger); Puglia: San Giovanni, Mte. Gargano, 15.5.1990 (leg. A. Müller);

Sardinia: Lanusei env., 29.6.2000 (leg. J. Halada); Brunella env., 70 m, 28.5.2013 (leg. J. Halada); Sicilia: 35 km

SW Ragusa, 18–22.6.2002 (leg. J. Halda). CROATIA: Buzet, 15.5.1993 (leg. Z. Pedr); Dvenik, 16.6.2000 (leg. M.

Kafka); 30 km SE Knin, 450 m, 25.5.2005 (leg. Z. Pedr); 40 km N Split, 370 m, 29.5.2005 (leg. Z. Pedr).

GREECE: Ionian Islands: Korfu, Mandouki, 19.5.1984 (leg. L. Norén); Epirus: 10 km E Parga, 14.6.1997 (leg. P.

Thomas); Thessaly: Mt. Ossa, Kokino Nero, 40 km NE Larissa, 13.5.2005 (leg. J. Halada); Central Greece: Euböä,

Xero mountains NE Drimona, 700 m, 26.5.2009 (leg. H. Rausch); Western Greece: Olympia, Alfios valley,

4.6.1995 (leg. W. Arens); Peloponnese: Mani, Agios Dimitrios, 25 m, 3.5.2001 (leg. A. Müller); Crete: Prina env.,

7 km S Istro, 12.6.2002 (leg. K. Denes); Aegean Islands: Lesbos, 2.7 km SSE Agiasos, 700 m, 21.6.2004 (leg. A.

Kyriakopoulos); Rhodos, Malonas, 150 m, 14.5.2005 (leg. A. Müller); Kos, 26.5.2008 (leg. G. Reder); Ikaria,

Glaredo, 20.4.2012 (leg. I. Vavitsas); Limnos, Phalakros, 13.5.2012 (J. Devalez); Naxos, Panagia Drosiani,

16.5.2012 (leg. I. Vavitsas); Thasos, Melissourgos, 22.5.2012 (leg. M. de Courcy); Samothrace, Alonia, 29.5.2012

(M. de Courcy); Chios, Aghio Gala, 7.6.2012 (P. Toutziarakis); Aegina, Sfentouri, 10.5.2013 (leg. S. Margaroni);

Santorini, Panagia Kalou, 11.5.2013 (leg. T. Petanidou); Kea, Dihala Sklavou, 26.5.2013 (leg. T. Petanidou).

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BULGARIA: Kresna, 5.1980 (leg. M. Kocourek); Nessebar, 29.6.1982 (leg. M. Kocourek); Rodopi, Hrabrino,

15.6.1997 (leg. A. Zaykov); Trakia, Proslav, 10.7.1997 (leg. Z. Pedr); Plovdiv, 15.8.1997 (leg. Z. Pedr); SE

Lozenec, 16.6.2008 (leg. M. Halada). TURKEY: Canakkale: Gelibou env., 18.6.1998 (leg. J. Halada); Izmir:

beween Ödmis and Bozdag, 1020 m, 28.5.2006 (leg. E. Scheuchl); Aydin: Kusadasi, 27.6.2006 (leg. E. Scheuchl);

Kütahya: 20 km NEE Kütahya, 13.7.2006 (leg. M. Halada); Burdur: around Dirimli mountainpass, 1350 m,

6.6.2006 (leg. E. Scheuchl); Antalya: Seklik Mevkli, 3.6.2009 (leg. J.S. Ascher, J. Rozen, H. Özbek); Ankara:

between Peçenek and Camlidere, 1140 m, 17.6.2006 (leg. E. Scheuchl); Aksaray: Ihara valley, 13.6.2008 (leg. M.

Kafka); Mersin: Kuzucubelen, 28.5.1998 (leg. M. Halada); Nevsehir: Göreme, 18.5.1988 (leg. N. Mohr); Adana:

Seyhan, 110 m, 22.5.2005 (leg. E. Scheuchl); Sivas: 45 km E Yarhisar, 24.6.1993 (leg. Jirousek); Hatay: Yayladagi,

11.6.1998 (leg. M. Halada); Malatya: Erkenek, 80 km SW Malatya, 9.7.1997 (leg. M. Halada); Adiyaman: E

Fidanlik, 800 m, 12.5.2006 (leg. H. Özbek); Sanliurfa: Caylarbasi, 70 km N Urfa, 2.6.1998 (leg. M. Halada);

Erzurum: Erzurum, Oltu Basakli, 1850 m, 5.6.2008 (leg. H. Özbek); Bitlis: 50 km E Tatvan, 7.7.1997 (leg. M.

Halada); Kars: Pasli, 50 km S Kars, 1.7.1997 (leg. M. Halada); Agri: Agri env., 27.6.1993 (leg. M. Halada).

CYPRUS: between Coral Bay and Agios Georgios, 15 km NNW Paphos, 5.5.2003 (leg. E. Scheuchl); Mesaoria,

Gaziköy W Vadili, 100 m, 26.4.2011 (leg. C. Sedivy, A. Müller); Karpaz peninsula, 29.4.2011 (leg. C. Sedivy, A.

Müller). ARMENIA: Aragatsotn: Parpi, 1600 m, 31.5.1973 (leg. A. Svozil); Mt. Aragats south slope, 3 km N

Byurakan, 1830 m, 22.6.2006 (leg. Ø. Berg); Yerevan: Yerevan, 1250 m, 12.6.2004 (leg. Ø. Berg). GEORGIA:

Ghvevi, 30 km W Tbilisi, 13.6.2015 (leg. M. Snizek). IRAN: Gilan: 5 km E Rudbar, 400 m, 8.6.2014 (leg. J.

Halada). TURKMENISTAN: Firyuza, 40 km W Aschabat, 6.6.1993 (leg. J. Halada). SYRIA: Marbij env., 9.5.1996

(leg. M. Halada); Jisr ash Shughur, 26.5.1996 (leg. M. Halada); Hamah/Masyaf env., 9.6.2000 (leg. K. Denes); 50

km W Homs, 12.5.2002 (leg. M. Halada). ISRAEL AND PALESTINE: Northern District: Golan heights, En Fit,

2.9 km ESE Senir, 4.5.2013 (A. Dorchin, D. Bénon, V. Trunz); Haifa District: 10 km S Haifa, Har Karmel/Bet

Oren, 14.5.1996 (leg. C. Schmid-Egger); Central District: 0.75 km N Yaqum, 12m, 9.4.2010 (leg. A. Dorchin); Tel

Aviv District: Tel Aviv, Botanical Garden, 5.4.2013 (leg. J.S. Ascher); Jerusalem District: Judean foothills, Park

Britannia, 24.4.2011 (leg. T. Koznichki); Southern District: 3 km N Lakhish, 1.4.2016 (leg. G. Pisanty). JORDAN:

Tall Al Arbatin, 20 km S North Shuna, 19.4.1996 (leg. M. Halada); 20 km NNW Al Karak, 27.4.2005 (leg. F.

Kantner); Dibbin, 15 km W Jerash, 2.5.2006 (leg. K. Denes); Wadi al Hasa S Al Karak, 20.4.2007 (leg. C. Praz, C.

Sedivy, A. Müller).

Distribution. From northernmost Morocco over Iberia (Portugal, Spain, Balearic Islands) and southern France

(including Corsica) to southwesternmost Switzerland; from eastern Austria, southern Slovakia and Hungary to

Romania; from Italy (including Sardinia and Sicily), Slovenia and the Balkan peninsula (Croatia, Montenegro,

Macedonia, Albania, Greece including Crete, Bulgaria) over Turkey and Cyprus to the Caucasus (Armenia,

Georgia), central Iran and southwestern Turkmenistan; from Turkey southwards to the Levant (Syria, Israel and

Palestine, Jordan). The westernmost record is from Sobreda near Lisbon in western Portugal, the northernmost

from Turňa nad Bodvou in southeastern Slovakia, the southernmost from Nurabad in Iranian Fars province and the

easternmost from Firyuza in southwestern Turkmenistan.

Pollen hosts. Oligolectic on Asteraceae (based on 50 pollen loads from 43 different localities in Bulgaria,

Cyprus, Greece, Israel and Palestine, Italy, Jordan, Morocco, Portugal, Spain, Turkey and Turkmenistan and on

field observations). Among the Asteraceae, O. ligurica only exploits the flowers of Asteroideae and Cichorioideae

with a preference for the former subfamily: 36 pollen loads were pure loads of Asteroideae pollen and four were

pure loads of Cichorioideae pollen, while seven were mixed loads consisting of the pollen of both subfamilies.

Three pollen loads contained small amounts of Brassicaceae, Campanulaceae or Resedaceae pollen beside

Asteroideae pollen, suggesting that the females very rarely also exploit pollen hosts other than the Asteraceae. On

the Asteroideae, the females take up pollen from the surface of the capitulum directly into their scopa by rapidly

moving the metasoma up and down (“abdominal drumming” sensu Cane 2017). Flower records: Hieracium

bauhinii (Tkalců 1994); Achillea spec., Anthemis tinctoria, Buphthalmum salicifolium, Calendula arvensis, Crepis

aculeata, Glebionis coronaria, Hypochaeris achyrophorus, Inula spec., Leontodon spec., Pallenis spinosa,

Pulicaria dysenterica, Reichardia picroides (label records).

Nesting biology. The nests are built in 3–4 mm wide burrows in the pith of dead plant stems excavated by

other aculeates (e.g. Ceratina) or in dead hollow stems (Friese 1893; Benoist 1931; Mavromoustakis 1939; Grandi

1961; G. Le Goff personal communication; A. Müller unpublished data; Fig. 9, 10). The most important nesting

sites are dead tendrils of Rubus; further nesting sites include dead stems of Artemisia, Foeniculum and Scolymus.

The nests contain up to eight linearly arranged brood cells, which are separated from each other by partitions

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OSMIINE BEES OF OSMIA (HOPLOSMIA)

constructed from leaf pulp. Leaf pulp is also used to seal the nest at its opening. Leucospis dorsigera Fabricius

(Leucospidae) was reared from the nests of O. ligurica (Baur & Amiet 2000).

Note. In the male, the lower half of the paraocular area is covered with a very short, appressed and plumose

pilosity. This specialized pilosity is hypothesized to be involved in the mating behaviour as it is often soaked with a

fluid of unknown origin, which might serve to mark home ranges or territories or to communicate with the female

during courtship and mating.

Osmia (Hoplosmia) olgae (Tkalců 1978)

Anthocopa (Odontanthocopa) olgae Tkalců 1978: 159. Type material: Holotype ♂, “Bulgaria sept.-or. Albena, 4.7.1976”

(Bulgaria), Oberösterreichisches Landesmuseum Linz.

Literature records. BULGARIA: Varna, between Varna and Vinica, between Balcik and Tuzlata, Balcik (Tkalců

1978). TURKEY: Nevsehir: Ürgüp (Özbek & Zanden 1992); 5km NW Ürgüp (Zanden 1996). The record of

Zanden (1996) from “Syria, Mts. Armatus” most probably refers to the Amanus mountains (= Nur mountains),

which formerly belonged to Syria but are now part of the Hatay province in southernmost Turkey.

New records. TURKEY: Kütahya: 20 km NNE Kütahya, 13.7.2006 (leg. M. Halada); Isparta: Karakus Dagi

centr., 1460 m, 11.7.2006 (leg. J. Halada); Karaman: Madensehir, 7.8.1972 (leg. K. Warncke); Nigde: Camardi,

13.7.1997 (leg. M. Halada); Nevsehir: 50 km E Ürgüp, 1100 m, 24.8.1991 (leg. K. Warncke); Ürgüp, 11.7.1997

(leg. M. Halada); Erzincan: 15 km W Refahye, W Erzincan, 1600 m, 7.7.2000 (leg. M. Halada); Artvin: Demirkent,

Yusufeli, 1650 m, 14.8.1991 (leg. E. Yildirim); Bitlis: Tatvan, 1720 m, 16.8.1991 (leg. K. Warncke).

Distribution. From the Black Sea coast of Bulgaria to eastern and southern Turkey.

Pollen hosts. The only two pollen loads available (from two different localities) consisted of pollen of

Asteroideae. Flower records: Centaurea arenaria, Melilotus albus (Tkalců 1978).

Nesting biology. Unknown.

Osmia (Hoplosmia) padri (Tkalců 1974)

Anthocopa (Odontanthocopa) padri Tkalců 1974a: 129. Type material: Holotype ♀, “Slancev Brjag, 28.6.–14.7.1971”

(Bulgaria), Pádr Collection Praha.

Literature records. CROATIA: Rijeka, Krk, Split (Zanden 1983); Barić Draga (Józan 2009). MONTENEGRO:

Ulcinj, Crna Gora/Budva Becici (Tkalců 1974a). MACEDONIA: Dojran Lake (Tkalců 1974a). GREECE:

Thessaly: Portaria/Pelionpass; Peloponnese: Epidaurus, Mykene (Zanden 1983). BULGARIA: Aitos, Arkutino,

Zadar-Sukosan, Sandanski (Tkalců 1974a); Albena, 17 km W Varna, Sandanski, Liljanovo (Tkalců 1979).

New records. CROATIA: Istria, Vodnjan env., 7.7.1999 (leg. Z. Pedr); Makarska, 27–30.6.2011 (leg. M.

Kadlecova). GREECE: Central Macedonia: Litochoro, Plaka, 10–28.7.1988 (leg. S. Risch); Attica: Sounion,

15.6.1966 (leg. W. Schläfle); Western Greece: Samikon, 23.7.1997 (leg. W. Arens); Michas–Erymanthos

mountains, 1300–1700 m, 27.7.1997 (leg. W. Arens); NW Niforeika-Kato, Achaia env., SW Patras, 3–17.7.2005

(leg. P. Bulirsch); Achaia near Chekali, 6.7.2006 (leg. W. Arens); Peloponnese: Mavromati, Ithome, 26.6.1996

(leg. W. Arens); Kalamata, Avia, 26.6.1996 (leg. W. Arens); Atsiholos, Gortis, 19.7.1997 (leg. W. Arens); Hosiari,

Ageranos, 1.7.1997 (leg. W. Arens); Lira, Lakonia, 3.7.1997 (leg. W. Arens); Arkadia, Orchomenos, 2.7.2001 (leg.

W. A r ens); Aegean Islands: Lesbos, 1.6 km NW Parakoila, 100 m, 23.6.2004 (leg. O. Messinger); Thasos, Potos,

17.6.2012 (leg. M. de Courcy). BULGARIA: Sandanski, 7.1967 (leg. M. Kocourek); Kresna-Pirin, 15.8.1982 (leg.

M. Kocourek); Primorsko env., 4–6.7.1988 (leg. P. Tyrner); SW Melnik, 13.8.1993 (leg. M. Halada); Stara Zagora,

5.7.2000 (leg. M. Snizek); SW Kresna, 150 m, 24.6.2008 (leg. M. Halada). TURKEY: Canakkale: 6 km N Ezine,

35 m, 27.06.2006 (leg. J. Halada); Aydin: Aydin, 4.7.2006 (leg. E. Scheuchl); Manisa: 40 km NW Salihli, 150 m,

26.6.2006 (leg. J. Halada); 30 km SEE Salihli, 430m, 29.6.2006 (leg. J. Halada); Denizli: 35 km SSE Denizli, 970

m, 5.7.2006 (leg. J. Halada); Burdur: 20 km SW Burdur, 1350 m, 8.7.2006 (leg. M. Halada); Antalya: 20 km E

Alanya, 16.6.1997 (leg. M. Halada); Isparta: 6 km E Egirdir, 15.7.1998 (leg. C. Schmid-Egger); 8 km NE Isparta,

1020 m, 9.7.2006 (leg. J. Halada); Egidir Gölü, 5 km N Akkecili, 920 m, 10.07.2006 (leg. J. Halada); Nevsehir: 10

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320

km W Ürgüp, 15.6.1998 (leg. M. Halada); Adana: Aladag, 780 m, 2.7.1995 (leg. P. Rasmont); Feke env., 21–

24.7.2000 (leg. M. Oboril); Sivas: 45 km E Yarhisar, 24.6.1993 (leg. M. Halada); Malatya: 15 km E Malatya,

27.6.2000 (leg. M. Halada); Elazig: SE Elazig, Hazar Gölü, 29.6.2000 (leg. M. Halada); Erzincan: Avcilar, 1250 m,

4.8.2003 (leg. S. Coruh); Mardin: 20 km N Mardin, 21.6.1997 (leg. M. Halada). SYRIA: Damascus, 22.5.1996

(leg. M. Halada); 40 km NE Damascus, 22.5.1996 (leg. M. Halada).

Distribution: From the Balkan Peninsula (Croatia, Montenegro, Macedonia, Greece, Bulgaria) eastwards to

eastern Turkey and from Turkey southwards to southern Syria.

Pollen hosts. Oligolectic on Asteraceae (based on 30 pollen samples from 22 different localities in Croatia,

Greece, Turkey and Syria). As all pollen loads analysed were pure loads of Centaurea, O. padri is likely a

specialist of Carduoideae. Flower records: Carduus spec. (Zanden, 1983), Centaurea arenaria (Tkalců 1979),

Centaurea solstitialis (Tkalců 1979, label record).

Nesting biology. The nests are built in dead wood (Tkalců 1974a), most probably in preexisting insect

burrows.

Osmia (Hoplosmia) scutellaris Morawitz 1868

Osmia scutellaris Morawitz 1868: 151. Type material: Syntypes ♂♀, “im Magnan-Thale” (France), “auch nördlicher bei

Pallanza” (Italy), “und Lugano” (Switzerland), type depository unknown.

Heriades integra Benoist 1934: 159. Type material: Holotype ♂, “Tanger” (Morocco). Muséum National d’Histoire Naturelle

Paris. Synonymy in Müller & Trunz (2014).

Literature records. ALGERIA: Bouira (Tkalců 1974a). SPAIN: Lérida (Ceballos 1956). ANDORRA: Andorra

(Zanden 1984b). FRANCE: Puy-de-Dôme, Drôme, Alpes-Maritimes, Var (Benoist 1931); Rhône, Vaucluse, Var,

Bouches-du-Rhône (Tkalců 1974a). SWITZERLAND: Valais, Ticino (Amiet et al. 2004). ITALY: Lazio,

Campania (Tkalců 1974a); Piemonte, Liguria, Friuli-Venezia Giulia, Emilia-Romagna, Umbria, Lazio, Abruzzi,

Molise, Campania, Basilicata (Pagliano 1994). SLOVAKIA: Kováčov near Štúrovo (Tkalců 1974a); Straka et al.

(2007). HUNGARY: Transdanubian Mountains (Józan 2011). UKRAINE: Crimea (Romasenko 1995). RUSSIA:

Dagestan/Derbent (Morawitz 1874); southern European Russia (Osychnyuk et al. 1978). SLOVENIA:

Submediterranean and subpannonian region (Gogala 1999). CROATIA: Brovinje, Cavići (Zagorje), Hreljin, Krk,

Lovran, Martina, Ravni, Sveta Jelena (Józan 2009). MACEDONIA: Ohrid, Katlanovska Banja (Zanden 1984a).

ALBANIA: Kula Ljums (Maidl 1922); Lukova (Tkalců 1974b). BULGARIA: Slantschev Brjag (Tkalců 1974a).

TURKEY: Icel (Özbek & Zanden 1992). IRAN: Tehran, Gilan and Mazandaran provinces (Nadimi et al. 2013).

New records. MOROCCO: Oriental: Taforalt, 660 m, 9.5.2002 (leg. H.J. Flügel); Fès-Meknès: SW Sefrou,

16.5.2003 (leg. J. Halada); Rabat-Salé-Kénitra: Pilote near Khemisset, 19.5.1997 (leg. Prudek); Souss-Massa:

Imouzzer NNW Agadir, 11.4.1988 (leg. J. Gusenleitner); Taroudant, 18.4.1990 (leg. M. Halada); Tamzergoute, 10

km N Agadir, 8.4.2015 (leg. C. Schmid-Egger). TUNISIA: Jendouba: Tabarka, Khathairia, 15.5.1993 (leg. M.

Hauser); 17 km N Tabarka, 16.4.2001 (leg. M. Snizek). PORTUGAL: Faro: Hortas do Tabual, 5 km SE Vila do

Bispo, 18.5.1995 (leg. J. Gusenleitner). SPAIN: Girona: Puigcerda, 18.7.1963 (leg. H. Hamann); Barcelona:

Papiol, 15.5.2010 (leg. J. Bosch); Tarragona: Montroig-Badia, 5 km WSW Cambrils, 14.5.2003 (leg. J. Smith);

Castellon: Castellon env., 6.1997 (leg. Kadlecova); Granada: Atalbeitar, 1250 m, 16.5.2014 (leg. J. Smith);

Malaga: Parauta, 13 km S Ronda, 800 m, 10.8.2004 (leg. S. Risch); Cadiz: Puerto Real, 19.4.2012 (leg. J. Smith).

FRANCE, Drôme: St. Thomé, 12 km SW Montélimar, 21.5.2000 (leg. J. van der Smissen); Va u c l u s e : Le Beaucet-

Venasque, 10 km SE Carpentras, 1.6.2000 (leg. J.van der Smissen); Var : Gonfaron, les Ribas, 180 m, 13.5.1997

(leg. Flagothier); Hérault: Marseillan, Agde, 7.5.1994 (leg. W. Arens); Aude: Carcassonne, Caunes-Minervois,

14.7.2002 (leg. J. Smith); Pyrénées-Orientales: 25 km S Prades, 990m, 21.7.2011 (leg. J. Halada); Corsica: Zonza,

19.6.1996 (leg. K. Denes). SWITZERLAND: Va l a i s : Visperterminen, Niederhäusern, 2.8.1997 (leg. W. Arens).

ROMANIA: Orsova, Svinita, 23.5.2002 (leg. J. Halada); Cozla, W of Turnu Severin, 26.5.2002 (leg. J. Halada).

UKRAINE: Krya, Jalta, Crimea, 8–10.7.1985 (leg. K. Denes); Stany env., Crimea, 27.5–2.6.2003 (leg. Andreeva).

ITALY: Liguria: Alassio, 7.7.1996 (leg. J. Tiefenthaler); Tos c a n a : Canneto, 25 km SSW Volterra, 13.7.1997 (leg. S.

Risch); Umbria: Aquasparta, Lo Scoppi, 5.6.1992 (leg. G. Zinnert); Lazio: Albano, 1.6.1987 (leg. G.G.M.

Schulten); Puglia: Mte. Gargano, San Giovanni, 5.7.1994 (leg. A. Müller); Sicily: Cammarata, 60 km N Agrigento,

1.6.2002 (leg. J. Halada). CROATIA: 30 km SE Knin, 450 m, 24.5.2005 (leg. Z. Pedr); 20 km W Neum, 25.5.2005

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OSMIINE BEES OF OSMIA (HOPLOSMIA)

(leg. Z. Pedr). SERBIA: Bieb Polje, 10 km NE Petrovac, 750 m, 19.6.2008 (leg. P. Banar). MACEDONIA: Dorjan,

4.6.1973 (leg. M. Kocourek); Galicica National Park, 600–1500 m, 26.6.2014 (leg. J. Halada, M. Fabianová).

GREECE: Ionian Islands: Corfu, Linia, 10.5.1984 (leg. L. Norén); Epirus: Ori Pargas, Sivota, 5.2000 (leg. F.

Kantner); Thessaly: 40 km NE Larissa, Mt. Ossa, Kokino Nero, 13.5.2005 (leg. M. Kadlecova); Western Greece:

Olympia, Alfios valley, 4.6.1995 (leg. W. Arens); Central Greece: Xerovouni-Geb., SE Pendagi, 1100 m, 8.6.2005

(leg. H. Rausch); Peloponnese: Mani, Agios Dimitrios, 20 m, 18.4.2001 (leg. A. Müller); Crete: Sougia, 8.4.2002

(leg. A. Müller); Aegean Islands: Chios, Diefcha, 16.5.2012 (leg. P. Toutziarakis); Ikaria, Glaredo, 28.5.2012 (leg.

I. Vavitsas); Karpathos, Olympos, 2.5.2012 (leg. I. Vavitsas); Lesbos, 2 km SSE Agiasos, 600m, 21.6.2004 (leg. A.

Kyriakopoulos); Naxos, Moutsouna, 20.5.2012 (leg. I. Vavitsas); Rhodos, Stegna, 20 m, 4.5.2005 (leg. A. Müller);

Samothrace, Alonia, 30.5.2012 (M. de Courcy); Thasos, Melissourgos, 23.5.2012 (M. de Courcy). BULGARIA:

Nessebar, 3.6.1982 (leg. M. Kocourek); Rodopi, Hrabrino, 5.7.1997 (leg. Z. Pedr); Trakia, Proslav, 10.7.1997 (leg.

Z. Pedr); Harmandli, 200 m, 14.6.2008 (leg. M. Halada). TURKEY: Canakkale: Canakkale, 9.6.1991 (leg. M.

Kocourek); Aydin : between Cayir and Ödemis, 705 m, 28.5.2006 (leg. E. Scheuchl); Kütahya: Porsuk Baraji, 30

km N Kütahya, 22.5.1998 (leg. M. Halada); Isparta: Dere mahallesi, 1150 m, 24.5.2004 (leg. H. Özbek); Antalya:

60 km E Antalya, Side-Kumköy, 4–17.4.2004 (leg. M. Herrmann); Kastamonu: between Toysa and Iskilip, 1170 m,

19.6.2006 (leg. E. Scheuchl); Ankara: between Kizlicahamam and Gerede, 1000 m, 17.6.2006 (leg. E. Scheuchl);

Konya: 10 km W Aksehir, 25.6.1998 (leg. J. Halada); Mersin: Aslanli, 30 km N Erdemli, 17.6.1998 (leg. M.

Halada); Nevsehir: Ürgüp, 30 km E Nevsehir, 1400 m, 30.5.2001 (leg. K. Denes); Hatay: Samandagi, 20.5.1995

(leg. K. Denes); Adiyaman: Kuyucak env., 10.6.2001 (leg. M. Snizek); Sanliurfa: 80 km E Birecik, 2.5.1995 (leg.

K. Denes); Artvin: Yusufeli Darica, 895 m, 16.6.2010 (leg. H. Özbek); Erzurum: Camlibel, Oltu, 1600 m,

26.6.2000 (leg. H. Özbek); Bitlis: Tatvan, 30.6.1993 (leg. K. Denes). CYPRUS: Paphos, Kato Paphos, 16.5.2009

(leg. D. Dauber); Karpaz Peninsula, 29.4.2011 (leg. C. Sedivy, A. Müller). GEORGIA: Ghvevi, 30 km W Tbilisi,

13.6.2015 (leg. M. Snizek). IRAN: Mazandaran: 10 km N Gashar, 2300–2700 m, 7.6.2014 (leg. J. Halada); Gilan:

5 km E Rudbar, 400 m, 8.6.2014 (leg. J. Halada). SYRIA: Jisr ash Shughur, 26.5.1996 (leg. M. Halada); 50 km W

Homs, 12.5.2002 (leg. M. Halada); Daraa, Wadi Al Marir, 20.5.1995 (leg. V. Nemec). ISRAEL AND PALESTINE:

Northern District: Golan, 2 km W Mas'ada, 900 m, 5.5.2010 (leg. C. Sedivy, C. Praz); Haifa District: 10 km S

Haifa, Har Karmel/Bet Oren, 14.5.1996 (leg. O. Niehuis); Central District: Beit Hanan, 26.4.2009 (leg. A.

Dorchin); Tel Aviv District: Tel Aviv, Botanical Garden, 5.4.2013 (leg. J.S. Ascher); Jerusalem District: Judean

foothills, Ya'ar Yish'I, 26.4.2011 (leg. T. Koznichki); Southern District: Judean Foothills, Beit Nir, 28.3.2010 (leg.

G. Pisanty). JORDAN: Aljun, 5.5.1995 (leg. K. Denes); Jordan Valey, Dayr Alla, 27.4.1996 (leg. M. Halada);

Saham, Irbid reg., 300 m, 19.4.2003 (leg. I. Plijushtch); Pella, 29.4.2007 (leg. K. Denes).

Distribution. From the Maghreb (Morocco, Algeria, Tunisia) over the Iberian Peninsula (Portugal, Spain,

Andorra) and southern France (including Corsica) to southern Switzerland; from southernmost Slovakia and

Hungary over Romania and southernmost Ukraine to southern European Russia; from Italy (including Sicily),

Slovenia and the Balkan Peninsula (Croatia, Serbia, Macedonia, Albania, Greece including Crete, Bulgaria) over

Turkey and Cyprus to the Caucasus (Georgia) and northern Iran; from Turkey southwards to the Levant (Syria,

Israel and Palestine, Jordan). The westernmost record is from Agadir in western Morocco, the northernmost from

Kováčov near Štúrovo in southernmost Slovakia, the southernmost from Beit Nir in Central Israel and the

easternmost from near Chalus in Iranian Mazandaran province.

Pollen hosts. Oligolectic on Asteraceae (based on 50 pollen loads from 42 different localities in Cyprus,

France, Greece, Israel and Palestine, Italy, Jordan, Morocco, Romania, Syria and Turkey and on field

observations). Among the Asteraceae, O. scutellaris only exploits species of Asteroideae and Cichorioideae with a

preference for the latter subfamily: 35 pollen loads were pure loads of Cichorioideae pollen and eight were pure

loads of Asteroideae pollen, while seven were mixed loads consisting of the pollen of both subfamilies. On the

Asteroideae, the females take up pollen from the surface of the capitulum directly into their scopa by rapidly

moving the metasoma up and down (“abdominal drumming” sensu Cane 2017). Flower records: Anthemis chia,

Chrysanthemum coronarium, Crepis aculeata, Inula spec., Pallenis spinosa, Picris spec., Sonchus asper, Thrincia

spec., Urospermum picroides (label records).

Nesting biology. The nests are built in burrows in the pith of dead plant stems excavated by other aculeates

(e.g. Ceratina) or in dead hollow stems (Friese 1893; Graeffe 1902; Benoist 1931; A. Gogala personal

communication). The most important nesting sites are dead tendrils of Rubus. The material to construct cell

partitions and nest plugs is still unknown, but is probably leaf pulp as in closely related species such as O. ligurica.

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Osmia (Hoplosmia) warnckei (Tkalců 1992)

Hoplosmia (Odontanthocopa) warnckei Tkalců 1992: 223. Type material: Holotype ♂, “15 km SE Sarvestan/Fars, 1800 m,

17.5.1978” (Iran), Oberösterreichisches Landesmuseum Linz.

Diagnosis of the hitherto unknown ♀: Osmia warnckei belongs to those O. (Hoplosmia) species, which have

their metanotum not hidden by the scutellum when seen from above and whose proboscis is not longer than the

mesosoma. Among these species, the female of O. warnckei is easily recognisable by the very coarse punctation of

head and mesosoma, the very sharp preooccipital margin and the comparably long proboscis, which is of about the

same length as the mesosoma (see Key to species).

Literature records. TURKEY: Sirnak: E of Sirnak, 5.6.1977 (Tkalců 1992).

New records. IRAN: Fars: Tangetamoradi near Yasouj, 4.6.2010 (leg. A. Monfared); Kalus near Yasouj,

30.5.2009 (leg. C. Sedivy, C. Praz, A. Monfared); Lorestan: 10 km SW Dorud, 1520m, 27.5.2014 (leg. J. Halada).

Distribution. From southeastern Turkey to central Iran.

Pollen hosts. The only pollen load available consisted of pollen of Centaurea (Asteraceae, Carduoideae).

Nesting biology. Unknown.

Key to species

The females of O. hermonensis and O. tyrneri are still unknown.

Females

1 Metanotum hidden by scutellum when seen from above; when seen in profile, posterior margin of scutellum either reaching

same level as metanotum or overhanging it. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

- Metanotum not hidden by scutellum when seen from above; when seen in profile, posterior margin of scutellum not reaching

same level as metanotum. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

2(1) Terga 1–3 red. First segm ent of labial palpus of roughly the same length as second segment, sometimes slightly longer or

shorter. Basitarsus of hind leg about 2.25x as long as its maximal width. (Body length 6.5–8 mm; Maghreb, Spain.) . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Osmia fallax Pérez 1895

- Terga black. First segment of labial palpus much shorter than second segment. Basitarsus of hind leg at least 2.75x as long as

broad . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3

3(2) Metasomal scopa black. (Body length 6–8 mm; Italy.). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Osmia picena (Tkalců 1999)

- Metasomal scopa yellowish or reddish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4(3) Posterior margin of scutellum rounded, reaching in profile not beyond metanotum. First section of posterior vein of second

submarginal cell shorter than third section (posterior vein is divided into three sections by the two recurrent veins). . . . . . . . . 5

- Posterior margin of scutellum flattened, reaching in profile beyond metanotum. First section of posterior vein of second sub-

marginal cell longer than or as long as third section. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

5(4) Pilosity of head, mesosoma and terga including scopa yellowish-brown (Fig. 11). Apical tergal hair bands whitish, appressed

and only slightly surpassing tergal margins (Fig. 11). Pilosity on tergal sides shorter: longest hairs on sides of terga 2 and 3 dis-

tinctly shorter than maximal width of marginal cell of forewing. Tegulae brownish to blackish. (Body length 7.5–10 mm;

Egypt, Levant.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Osmia carbo (Zanden 1994)

- Pilosity of head, mesosoma and terga including scopa bright orange (Fig. 12). Apical tergal hair bands orange, suberect and

distinctly surpassing tergal margins (Fig. 12). Pilosity on tergal sides longer: longest hairs on sides of terga 2 and 3 about as

long as maximal width of marginal cell of forewing. Tegulae orange. (Body length 7.5–11 mm; Maghreb, Spain.) . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Osmia pinguis Pérez 1895

6(4) Distance between posterior margin of lateral ocellus and postocular tangent (= supraorbital line) shorter than the diameter of

lateral ocellus. Body length not exceeding 8 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

- Distance between posterior margin of lateral ocellus and postocular tangent (= supraorbital line) as long as or exceeding the

diameter of lateral ocellus. Body length often exceeding 8 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

7(6) Scutum except marginal areas nearly hairless with very short hairs. Axillae covered with very short and dense yellowish-

brown pilosity. Punctation of propodeum beside propodeal pit very scattered, interspaces reaching the diameter of several

punctures. (Body length 5–6.5 mm; Italy, southeastern Europe, Turkey.) . . . . . . . . . . . . . . . . . . . . .Osmia croatica Friese 1893

- Scutum distinctly, albeit rather sparsely haired with long hairs. Axillae covered with long and sparse whitish pilosity. Puncta-

tion of propodeum beside propodeal pit dense, interspaces not exceeding the diam eter of one puncture. (Body length 6–8 mm;

Europe eastwards to Central Asia and eastern Siberia.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Osmia spinulosa (Kirby 1802)

8(6) Scutum and scutellum except marginal areas very sparsely haired, longest hairs on scutum less than 2x as long as the diameter

of one puncture. Pilosity of mesepisternum very sparse, not hiding the sculpture of the integument. Punctation of clypeus,

323

OSMIINE BEES OF OSMIA (HOPLOSMIA)

frons, vertex, thorax and terga coarse; punctation of supraclypeal area distinctly finer than punctation of basal half of clypeus.

(Body length 8–10 mm; Aegean Islands, Turkey, Jordan.). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Osmia elegans (Tkalců 1992)

- Scutum moderately densely and scutellum very densely haired, longest hairs on scutum about 5x as long as the diameter of one

puncture. Pilosity of mesepisternum dense, partly hiding the sculpture of the integument. Punctation of head, thorax and meta-

soma distinctly finer; punctation of supraclypeal area only slightly finer than punctation of basal half of clypeus . . . . . . . . . . 9

9(8) Preoccipital margin sharp but not distinctly raised (Fig. 18). Inner margin of apicalmost part of fore tibial spur slightly convex

(Fig. 20). Scopa yellowish-white (Fig. 16). Pilosity of vertex and upper side of mesosoma yellowish-white (Fig. 16). Anterior

side of antennal segments 7–11(12) usually more or less reddish-brown (Fig. 16). Impunctate zone of apical margins of terga

1–2 usually broader and often more or less reddish-brown. (Body length 7.5–9 mm; Levant.) . . . Osmia centaureae spec. nov.

- Preoccipital margin sharp and distinctly raised (Fig. 19). Inner margin of apicalmost part of fore tibial spur straight to concave

(Fig. 21). Scopa yellowish-red. Pilosity of vertex and upper side of mesosoma yellowish-brown. Antenna entirely dark (Fig.

19). Impunctate zone of apical margins of terga 1–2 usually narrower and always dark. (Body length 8–10 mm; southeastern

Europe, Turkey, Levant.). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Osmia spinigera Latreille 1811

10(1) Proboscis distinctly longer than mesosoma, second segment of labial palpus at least 3.75x as long as first segment. . . . . . . . 11

- Proboscis of about the same length as mesosoma or shorter, second segment of labial palpus at most 3x as long as first segment

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .14

11(10) Clypeus with narrow impunctate apical margin. Apical fourth of clypeus distinctly flattened, its punctation uniform and much

finer than that on median part of the clypeus.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .12

- Apical margin of clypeus punctured. Apical fourth of clypeus slightly bent and not flattened, its punctation irregular and com-

posed of both small and large punctures. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .13

12(11) Metasomal scopa yellowish. Scutum sparsely haired, longest hairs about 7–8x as long as the diameter of one puncture. Anten-

nal segm ents 4 and 5 nearly as long as wide. (Body length 7–9 mm.) No morphological characters are known to distinguish the

following two closely related species in the female sex. However, identification is possible due to their non-overlapping distri-

bution ranges: . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Osmia dido Gribodo 1894: Maghreb

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . … Osmia distinguenda (Tkalců 1974): Peloponnese, Turkey, Levant

- Metasomal scopa white. Scutum except marginal areas nearly hairless, longest hairs about 2–3x as long as the diameter of one

puncture. Antennal segments 4 and 5 distinctly shorter than wide. (Body length 6.5–7.5 mm; southeastern Europe, Turkey,

Syria.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Osmia padri (Tkalců 1974)

13(11) Apical margin of clypeus medially distinctly emarginate. Width of emarginated median part of apical clypeal margin less than

1.5x as wide as length of converging lateral part of clypeal margin. (Body length 8–10 mm; Maghreb, southw estern Europe.)

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Osmia anceyi Pérez 1879

- Apical margin of clypeus medially straight. Width of straight median part of apical clypeal margin more than 1.5x as wide as

length of converging lateral part of cly peal margin. (Body length 7.5–10 mm; Italy, southeastern and eastern Europe, Turkey,

Levant, Iran, Egypt.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Osmia bidentata Morawitz 1876

14(10) Proboscis about as long as mesosoma, reaching in repose to middle of fore coxa; second segment of labial palpus almost as

long as length of compound eye. Punctation of gena, frons, vertex and mesosoma very coarse; diameter of largest punctures on

vertex about half as long as ocellar diameter. Preoccipital margin very sharp. (Body length 8–8.5 mm; Turkey, Iran.). . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Osmia warnckei (Tkalců 1992)

- Proboscic distinctly shorter than mesosoma, reaching in repose not to middle of fore coxa; second segment of labial palpus dis-

tinctly shorter than length of compound eye. Punctation of gena, frons, vertex and mesosoma much finer, diameter of largest

punctures on vertex at most one fifth as long as ocellar diameter. Preoccipital margin less sharp. . . . . . . . . . . . . . . . . . . . . . . 15

15(14) Head slightly wider than long. Distance between lateral ocellus and preccipital ridge at least 2.5x as long as ocellar diameter.

Clypeus in profile almost flat; its medioapical part impressed with the lateral corners slightly raised. Tibial spurs of hind leg

yellowish-brown. (Body length 7–9 mm; Morocco, southern and eastern Europe, Turkey, Levant, Iran, Turkmenistan.) . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Osmia ligurica Morawitz 1868

- Head slightly longer than wide. Distance between lateral ocellus and preccipital ridge at most 2x as long as ocellar diameter.

Clypeus in profile arched; apical margin with two more or less distinct tubercles separated by a distinct impression. Tibial

spurs of hind leg yellowish-brown or blackish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .16

16(15) Pilosity on mesepisternum short, much shorter than the longest hairs on scutum. Maximal width of scutellum less than twice as

wide as maximal width of axilla. Longest hairs along inner side of hind tibia about half as long as maximal tibial width. Tibial

spurs of hind leg yellowish-brown (always?). (Body length 5.5–6 mm; Bulgaria, Turkey.) . . . . . . . Osmia olgae (Tkalců 1978)

- Pilosity on mesepisternum long and of about the same length as on scutum. Maximal width of scutellum at least twice as wide

as maximal width of axilla. Longest hairs along inner side of hind tibia distinctly longer than half of maximal tibial width. Tib-

ial spurs of hind leg blackish. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .17

17(16) Converging margins of apical part of labrum almost straight, narrowing into a small apex. Preoccipital margin rounded. Bend

of hypostomal carina abrupt, lateral branches straight. (Body length 6.5 mm; Canary Islands.) . . . Osmia larochei Tkalců 1993

- Converging margins of apical part of labrum broadly rounded, narrowing into a wide apex. Preoccipital margin sharp to cari-

nate. Bend of hypostomal carina broadly rounded, lateral branches curved. (Body length 6–8 mm; Maghreb, southern and east-

ern Europe, Turkey, Levant, Iran.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Osmia scutellaris Morawitz 1868

MÜLLER

324

Males

1 Terga 1–3 red. First segment of labial palpus of roughly the same length as second segment, sometimes slightly longer or

shorter. (Body length 6.5–8 mm; Maghreb, Spain.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Osmia fallax Pérez 1895

- Terga black. First segment of labial palpus much shorter than second segment. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2(1) Preapical margin of tergum 6 with long spines. Apical margin of tergum 7 with median tooth. . . . . . . . . . . . . . . . . . . . . . . . . . 3

- Preapical margin of tergum 6 without distinct spines, sometimes crenulate or denticulate. Apical margin of tergum 7 rounded,

emarginate or bifid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

3(2) Sternum 1 without spine. Body length at most 6.5 mm. (Body length 5–6.5 mm; Italy, southeastern Europe, Turkey.). . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Osmia croatica Friese 1893

- Sternum 1 with spine. Body length usually longer than 6.5 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4(3) Spine of sternum 1 pointed or rounded. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

- Spine of sternum 1 bifurcate or emarginate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7

5(4) Scutum and scutellum except marginal areas very sparsely haired, longest hairs less than 2x as long as the diameter of one

puncture. Mesepisternum sparsely covered with short, thick, distinctly plumose und mostly appressed hairs, in its upper half

almost hairless. Preapical margin of tergum 5 with distinct spines, which are partly hidden by the white apical hair band.

Antenna reddish-brown in its apical half. Body length often surpassing 8.5 mm. (Body length 8–10 mm; Aegean Islands, Tur-

key, Jordan.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Osmia elegans (Tkalců 1992)

- Scutum and scutellum rather densely covered with long hairs, which are four or more times longer than the diameter of one

puncture. Mesepisternum densely covered with long, thin, weakly plumose and erect hairs, in its upper half as densely haired

as in its lower half. Preapical margin of tergum 5 with indistinct short tubercles or very short spines. Antenna dark. Body

length not surpassing 8.5 mm.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6

6(5) Spine of sternum 1 narrowly triangular when seen from below, its lateral margins distinctly concave and almost parallel-sided

in the apical third. Punctation of terga distinctly finer and less dense, interspaces on terga 2–3 medially often reaching or even

surpassing the diameter of one puncture, on tergum 6 basally very narrow but distinct. Pilosity of vertex, scutum and scutellum

yellowish-brown in fresh specimens. Apex of gonoforceps sharply pointed (Tkalců 1992). (Body length 6.5–8 mm; Europe

eastwards to Central Asia and eastern Siberia.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Osmia spinulosa (Kirby 1802)

- Spine of sternum 1 broadly triangular when seen from below, its lateral margins almost straight and regularly tapering towards

the apex. Punctation of terga distinctly coarser and denser, interspaces on terga 2–3 medially rarely surpassing the diameter of

half a puncture, on tergum 6 basally almost lacking. Pilosity of vertex, scutum and scutellum greyish-white in fresh specimens.

Apex of gonoforceps less sharply pointed (Tkalců 1992). (Body length 8–8.5 mm; Central Asia.) . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Osmia tyrneri (Tkalců 1992)

7(4) Punctation of sterna 2–4 fine, shallow and scattered, interspaces often exceeding the diameter of three or more punctures.

Punctation of terga 1–4 fine and moderately dense with interspaces usually exceeding the diameter of one puncture. Distance

between lateral ocellus and preoccipital margin less than 2.5x as long as ocellar diameter. Preoccipital margin acutely angled

but not sharp. (Body length 7–8 mm; Italy.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Osmia picena (Tkalců 1999)

- Punctation of sterna 2–4 coarse, deep and rather dense, interspaces only rarely exceeding the diameter of two punctures. Punc-

tation of terga 1–4 coarse and dense with interspaces rarely reaching the diameter of one puncture. Distance between lateral

ocellus and preoccipital margin more than 2.5x as long as ocellar diameter. Preoccipital margin sharp. . . . . . . . . . . . . . . . . . .8

8(7) Preoccipital margin sharp but not distinctly raised. Apicalmost part of gonoforceps with many single erect hairs (Fig. 24).

Inner margin of apicalmost part of fore tibial spur straight to slightly convex. Antennal segments (5)6–13 more or less yellow-

ish-brown to orange. Densely shagreened basal part of tergum 6 with very weak and almost im perceivable punctation (Fig.

22). Apical margins of terga 1–3 usually more or less reddish–brown and apically often very weakly bent upwards. (Body

length 7–9.5 mm; Levant.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Osmia centaureae spec. nov.

- Preoccipital margin sharp and distinctly raised. Apicalmost part of gonoforceps with dense tuft of hairs directed alongside and

slightly surmounting the gonoforceps (Fig. 25). Inner margin of apicalmost part of fore tibial spur distinctly concave. Antenna

entirely dark, occasionally more or less dark reddish-brown in its apical half. Densely shagreened basal part of tergum 6 with

rather distinct albeit shallow punctation (Fig. 23). Apical margins of terga 1–3 dark and flat. (Body length 7.5–11 mm; south-

eastern Europe, Turkey, Levant.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Osmia spinigera Latreille 1811

9(2) Proboscis distinctly longer than mesosoma, second segment of labial palpus at least 3.75x as long as first segment. Apical mar-

gin of tergum 7 bifid to weakly emarginate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

- Proboscis of about the same length as mesosoma or shorter, second segment of labial palpus at most 3x as long as first seg-

ment. Apical margin of tergum 7 rounded to weakly emarginate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

10(9) Apex of gonoforceps rounded on the inside. Sternum 4 medially with inconspicuous narrow longitudinal zone of appressed

white hairs, its apical margin usually more or less distinctly thickened and rarely emarginate medially. . . . . . . . . . . . . . . . . . 11

- Apex of gonoforceps acutely angled on the inside. Sternum 4 medially without narrow longitudinal zone of appressed white

hairs, its apical margin usually not distinctly thickened and often weakly emarginate medially. . . . . . . . . . . . . . . . . . . . . . . .13

11(10) Apical margin of tergum 7 medially very weakly and shallowly emarginate. Hidden basal zone of terga 5–6 sharply impressed,

completely dull and with much finer punctation than on disc. (Body length 9–9.5 mm; Israel.). . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Osmia hermonensis (Tkalců 1992)

- Apical margin of tergum 7 bifid. Hidden basal zone of terga 5–6 weakly impressed and with similar sculpture and punctation

as on disc . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

12(11) Tergum 7 usually dark. Apical zone of sternum 4 usually dark. Gonoforceps apically with evenly rounded outer margin, its api-

calmost part less button-like. Dorsal surface of gonoforceps apically impunctate, rarely with one or two punctures. (Body

length 7.5–9 mm; Maghreb.). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Osmia dido Gribodo 1894

325

OSMIINE BEES OF OSMIA (HOPLOSMIA)

- Tergum 7 usually reddish-brown to orange. Apical zone of sternum 4 usually orange coloured. Gonoforceps apically with

almost straight outer margin, its apicalmost part button-like. Dorsal surface of gonoforceps apically punctured. (Body length

7–9.5 mm; Peloponnese, Turkey, Levant.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Osmia distinguenda (Tkalců 1974)

13(10) Distance between lateral ocellus and preoccipital margin more than 2.5x as long as ocellar diameter. Scutum sparsely haired,

longest hairs about 5x as long as the diameter of one puncture or longer. Antennal segment 4 longer than broad, distinctly lon-

ger than segment 3. Body length at least 7.5 mm. (Body length 7.5–9.5 mm.) No morphological characters are known to distin-

guish the following two closely related species in the male sex. However, identification is possible due to their non-

overlapping distribution ranges: . . . . . . . . . . . . . . . . . . . . . . . . . . . . Osmia anceyi Pérez 1879: Maghreb, southwestern Europe

. . . . . . . . . . . . . . . . . .… Osmia bidentata Morawitz 1876: Italy, southeastern and eastern Europe, Turkey, Levant, Iran, Egypt

- Distance between lateral ocellus and preoccipital ridge less than 2.5x as long as ocellar diameter. Scutum except marginal areas

nearly hairless, longest hairs about 2x as long as the diameter of one puncture. Antennal segment 4 quadrate, about as long as

segment 3. Body length at most 7.5 mm. (Body length 6.5–7.5 mm; southeastern Europe, Turkey, Syria.). . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Osmia padri (Tkalců 1974)

14(9) Metanotum almost vertical and hidden by scutellum when seen from above. Tergum 7 large, its apical margin broadly rounded.

Sternum 2 apically rounded and medially sparsely punctured with interspaces reaching the diameter of several punctures; its

apical fifth often slightly bent up and separated from the basal four fifth by a narrow transverse impression. Sterna 3 and 4 with

a narrow longitudinal row of short white hairs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

- Metanotum less steeply inclined and entirely visible when seen from above. Tergum 7 small, its apical margin rounded to

weakly emarginate. Sternum 2 apically straight and densely punctured throughout with interspaces rarely exceeding the diam-

eter of one puncture; its apical fifth unmodified. Sterna 3 and 4 without longitudinal rows of short white hairs.. . . . . . . . . . . 16

15(14) Pilosity of head, mesosoma and terga yellowish-brown. Apical tergal hair bands whitish, more or less appressed and short,

covering only the apical half of the marginal zone of terga 2–4 or less. Apical margin of sternum 2 usually with small button-

like median protuberance (Fig. 13). Bristle-beset zone on dorsolateral side of gonoforceps shorter: distance between tip of

gonoforceps and insertion of the most basal bristles shorter than the length of the bristles arising from the tip of the gonofor-

ceps (Fig. 14). (Body length 7.5–10mm; Egypt, Levant.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Osmia carbo (Zanden 1994)

- Pilosity of head, mesosoma and terga bright orange. Apical tergal hair bands orange, suberect and long, covering much of the

marginal zone of terga 2–4. Apical margin of sternum 2 usually without small button-like median protuberance. Bristle-beset

zone on dorsolateral side of gonoforceps longer: distance between tip of gonoforceps and insertion of the most basal bristles as

long as or longer than the length of the bristles arising from the tip of the gonoforceps (Fig. 15). (Body length 7.5–11mm;

Maghreb, Spain.). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Osmia pinguis Pérez 1895

16(14) Apical margin of tergum 7 weakly emarginate. Gonoforceps very broad, distinctly widened apically . . . . . . . . . . . . . . . . . . .17

- Apical margin of tergum 7 rounded, rarely truncated. Gonoforceps slender, more or less parallel-sided over its whole length. .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .18

17(16) Proboscis distinctly shorter than mesosoma, not reaching in repose to middle of fore coxa; second segment of labial palpus dis-

tinctly shorter than length of compound eye. Punctation of gena, frons, vertex and mesosoma much finer, diameter of largest

punctures on vertex at most one fifth of one ocellar diameter. Antennal segment 3 distinctly longer than segment 4. Lower half

of paraocular area covered with very short, appressed and plumose pilosity. Preoccipital margin less sharp. (Body length 7–8.5

mm; Morocco, southern and eastern Europe, Turkey, Levant, Iran, Turkmenistan.) . . . . . . . . . . Osmia ligurica Morawitz 1868

- Proboscis about as long as mesosoma, reaching in repose to middle of fore coxa; second segment of labial palpus almost as

long as length of compound eye. Punctation of gena, frons, vertex and mesosoma very coarse; diameter of largest punctures on

vertex about half as long as ocellar diameter. Antennal segment 3 about as long as segment 4. Lower half of paraocular area

covered with normal pilosity. Preoccipital margin very sharp. (Body length 8–8.5 mm; Turkey, Iran.) . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Osmia warnckei (Tkalců 1992)

18(16) Pilosity of mesepisternum short, much shorter than the longest hairs on scutum. Maximal width of scutellum less than twice as

wide as maximal width of axilla. Tergum 7 reddish-brown. (Body length 5–6 mm; Bulgaria, Turkey.). . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Osmia olgae (Tkalců 1978)

- Pilosity of mesepisternum long and of about the same length as on scutum. Maximal width of scutellum more than twice as

wide as maximal width of axilla. Tergum 7 dark.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

19(18) Tergum 6 preapically with two indistinct lobes, which are weakly separated from each other by a shallow impression and have

a finely crenulate apical margin. Converging margins of apical part of labrum almost straight, narrowing into a small apex. Pre-

occipital margin rounded. Apical fourth of sternum 4 distinctly bent. (Body length 5.5 mm; Canary Islands.) . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Osmia larochei Tkalců 1993

- Tergum 6 preapically with two prominent lobes, which are distinctly separated from each other by a deep impression and have

an entire to coarsely crenulate apical margin. Converging margins of apical part of labrum broadly rounded, narrowing into a

wide apex. Preoccipital margin sharp to carinate. Sternum 4 almost flat, its apical fourth only slightly bent. (Body length 5.5–

7.5 mm; Maghreb, southern and eastern Europe, Turkey, Levant, Iran.) . . . . . . . . . . . . . . . . . Osmia scutellaris Morawitz 1868

Acknowledgments

M. Schwarz (Ansfelden) and F. Gusenleitner (Oberösterreichisches Landesmuseum Linz) loaned material of O.

(Hoplosmia) for study. C. Praz (University of Neuchâtel) and C. Sedivy (Zurich) participated in excursions to

Jordan and Tunisia. E. Neubert (Natural History Museum Bern) helped with species identification of snail shells. G.

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326

Pagliano (University of Torino) provided information on the occurrence of Osmia anceyi and O. bidentata in Italy.

A. Gogala (Slovenian Museum of Natural History), G. Le Goff (Barentin), J. Ortiz-Sanchez (University of

Almeria), G. Pisanty (Tel Aviv University) and H. Wiesbauer (Wien) conveyed unpublished observations on the

nesting biology of O. (Hoplosmia) species. A. Gogala, G. Pisanty and H. Wiesbauer provided photos. H. Baur

(Natural History Museum Bern) provided access to a digital imaging system for taking photomicrographs.

Suggestions and comments by J. Gardner, J. Gibbs and an anonymous reviewer improved the manuscript.

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(Monograph) The wild bees (Hymenoptera: Apoidea) of Morocco

Article

Full-text available

Dec 2020

Morocco is a well known hot-spot of biodiversity in the Mediterranean basin. While some taxa like vascular plants are relatively well recorded, important groups of pollinators like bees are still understudied. This article presents an updated checklist of the bee species of Morocco and includes a summary of global and regional distribution of each species. A total of 961 species belonging to six bee families and 68 genera are recorded: Andrenidae (8 genera, 217 species); Apidae (15 genera, 241 species); Colletidae (2 genera, 74 species), Halictidae (12 genera, 144 species), Megachilidae (28 genera, 271 species) and Melittidae (3 genera, 14 species). Among them, 67 species are recorded for the first time in Morocco. Around 70% of the bee fauna of Morocco consists of widespread Palaearctic species. Only 18% of Moroccan species recorded are restricted to North Africa and 8% are Moroccan single-country endemics (81 species). Afrotropical elements in the Moroccan fauna are few, with only 3% of Morocco species co-occuring in that region. This checklist is intended to stimulate new regional research on bees including their taxonomy and biogeography. As many groups of bees have been understudied, discovery of new species for science and new records for the country can be expected. Additional research including inventorying, monitoring, and integrative taxonomic studies are needed to develop a comprehensive strategy for bee conservation in Morocco.

NEW AND INTERESTING DATA OF WILD BEES (HYMENOPTERA, APOIDEA, ANTHOPHILA) FROM THE CANTABRIAN AREA (NORTHERN SPAIN), INCLUDING A SPECIES NEWLY RECORDED FOR SPAIN AND THE CONFIRMATION OF THE PRESENCE OF NOMADA ERRANS LEPELETIER, 1841 ON THE IBERIAN PENINSULA

Article

Full-text available

Jun 2021

Data of Anthophila species that have not been previously recorded in the Cantabrian area of Spain are presented. Also, some first records for some provinces, confirmation of some taxa whose presence was previously seen as doubtful for various reasons, and some new records of species that are rare or had only rarely been detected in the study area are provided. Additionally, Hylaeus incongruus Förster, 1871 is recorded for the first time for Spain and the presence of Nomada errans Lepeletier, 1841 on the Iberian Peninsula is confirmed. Novedades faunísticas interesantes de abejas (Hymenoptera, Apoidea, Anthophila) de la cornisa cantábrica (norte de España), incluida una especie nunca registrada en España y la confirmación de la presencia de Nomada errans Lepeletier, 1841 en la península ibérica Resumen: Se presentan datos de especies de Anthophila nunca antes citadas en la cornisa cantábrica de España. Así mismo, se proporcionan primeras citas en algunas de las provincias, confirmación de ciertos taxones de presencia dudosa por diversas razones y algunas citas nuevas de especies raras o con escasos registros en el área de estudio. Además, se cita Hylaeus incongruus Förster, 1871 por primera vez en España y se confirma la presencia de Nomada errans Lepeletier, 1841 en la península ibérica.

National records of 3000 European bee and hoverfly species: A contribution to pollinator conservation

Article

Full-text available

Sep 2023
INSECT CONSERV DIVER

Pollinators play a crucial role in ecosystems globally, ensuring the seed production of most flowering plants. They are threatened by global changes and knowledge of their distribution at the national and continental levels is needed to implement efficient conservation actions, but this knowledge is still fragmented and/or difficult to access. As a step forward, we provide an updated list of around 3000 European bee and hoverfly species, reflecting their current distributional status at the national level (in the form of present, absent, regionally extinct, possibly extinct or non-native). This work was attainable by incorporating both published and unpublished data, as well as knowledge from a large set of taxonomists and ecologists in both groups. After providing the first National species lists for bees and hoverflies for many countries, we examine the current distributional patterns of these species and designate the countries with highest levels of species richness. We also show that many species are recorded in a single European country, highlighting the importance of articulating European and national conservation strategies. Finally, we discuss how the data provided here can be combined with future trait and Red List data to implement research that will further advance pollinator conservation.

Twenty New Records of Bees (Hymenoptera, Apoidea) for Sardinia (Italy)

Article

Full-text available

Jul 2021

In Sardinia, the second largest Mediterranean island, 316 species of bees are known. Here, for the first time, the following 20 taxa are reported: Colletes cunicularius (Linnaeus, 1761), and C. eous Morice, 1904 (Colletidae); Andrena humilis Imhoff, 1832, A. granulosa Pérez, 1902, A. cineraria (Linnaeus, 1758), A. pallitarsis Pérez, 1903, A. rugulosa Stöckhert, 1935, A. savignyi Spinola, 1838, and A. tenuistriata Pérez, 1895 (Andrenidae); Sphecodes reticulatus Thomson, 1870 (Halictidae); Lithurgus tibialis Morawitz, 1875, Chelostoma emarginatum (Nylander, 1856), Dioxys cinctus (Jurine, 1807), Coelioxys caudatus Spinola, 1838, C. obtusus Pérez, 1884, and Megachile ericetorum (Lepeletier, 1841) (Megachilidae); and Nomada melathoracica Imhoff, 1834, N. pulchra Arnold, 1888, Eucera proxima Morawitz, 1875 and Tetralonia malvae (Rossi, 1790) (Apidae). N. pulchra is reported for the first time in Italy.

Andrena species (Hymenoptera: Apoidea: Andrenidae) from Western Algeria, with a preliminary assessment of their pollen preferences

Article

Full-text available

Apr 2021

Algeria, the largest country in Africa, has a species rich bee fauna because of its ecological conditions, climate and diversified flora. The present work represents a list of the genus Andrena Fabricius, 1775 in western Algeria. The survey, carried out during the period 2017–2019, has allowed identification of 56 species of Andrena in a total of 787 specimens (496 females, 291 males), with one species new for the fauna of Algeria, Andrena (Aenandrena) hystrix Schmiedeknecht, 1883. Preliminary pollen preferences of the most common species were analysed in Algeria for the first time, comprising seven polylectic and 12 oligolectic species. The current work provides a baseline for future studies on the diet of Andrena and is one of the first studies on the floral preferences of North African bees.

Annotated checklist of the megachilid bees of Corsica (Hymenoptera, Megachilidae)

Article

Full-text available

Mar 2024
J HYMENOPT RES

Corsica stands as one of the largest Mediterranean Islands and has been the exploration ground for renowned entomologists like Charles Ferton. However, no synthesis on Corsican bees has been published so far. To fill this gap in knowledge, we propose an overview of the megachilid bee fauna of the island based on fieldwork, a thorough examination of material housed at the Muséum national d’Histoire naturelle (MNHN, Paris), data compilation from various collections and a comprehensive review of existing literature. We reviewed 5,886 specimens and we extracted 279 additional data from literature sources. These data confirm the presence in Corsica of 91 species of which two are endemic, including Hoplitis corsaria (Warncke, 1991) which is elevated to species rank stat. nov. One new synonymy is established: Megachile lucidifrons Ferton, syn. nov. of Megachile albocristata Smith, 1853. The presence in Corsica of 19 species is regarded as dubious or erroneous. Finally, the types of Megachilid bees housed at MNHN and described based on Corsican material are illustrated. Lectotypes are designated for Megachile sicula var. corsica Benoist, 1935, Osmia corsica Ferton, 1901, Osmia erythrogastra Ferton, 1905, Osmia lanosa Pérez, 1879, and Osmia lineola Pérez, 1895.

Contribution to the knowledge of the bee fauna (Hymenoptera, Apoidea, Anthophila) in Serbia

Article

Full-text available

Aug 2021

The current work represents summarised data on the bee fauna in Serbia from previous publications, collections, and field data in the period from 1890 to 2020. A total of 706 species from all six of the globally widespread bee families is recorded; of the total number of recorded species, 314 have been confirmed by determination, while 392 species are from published data. Fourteen species, collected in the last three years, are the first published records of these taxa from Serbia: Andrena barbareae (Panzer, 1805), A. clarkella (Kirby, 1802), A. fulvicornis (Schenck, 1853), A. intermedia (Thomson, 1870), A. lapponica (Zetterstedt, 1838), A. pandellei (Pérez, 1895), A. paucisquama (Noskiewicz, 1924), A. simillima (Smith, 1851), Panurginus herzi (Morawitz, 1892), Epeoloides coecutiens (Fabricius, 1775), Nomada leucophthalma (Kirby, 1802), Chelostoma nasutum (Pérez, 1895), Hoplitis claviventris (Thomson, 1872), and Dasypoda pyrotrichia (Förster, 1855). Almost all the species recorded so far in Serbia belong to the West-Palaearctic biogeographical region, except Megachile sculpturalis (Smith, 1853), which is an alien invasive species native to East Asia. According to the European Red List of bees, 221 species listed in this paper were assessed as Data Deficient; threatened species mostly belong to the families Apidae with 13 species, Colletidae with eight species, and Halictidae with five species. This study contributes to the knowledge of the distribution of bee species in Europe. The present work provides a baseline for future research of wild bee diversity in Serbia and neighbouring regions at the local and regional levels, and a basis for their conservation.

The bees of Lebanon (Hymenoptera: Apoidea: Anthophila)

Article

Full-text available

May 2021

The study of wild bees has markedly increased in recent years due to their importance as pollinators of crops and wild plants, and this interest has been accentuated by increasing evidence of global declines in their abundance and species richness. Though best studied in Europe and North America, knowledge on the current state of wild bees is scarce in regions where they are particularly diversified, such as the Mediterranean basin. The eastern Mediterranean country of Lebanon, located at the heart of the Levant in a biodiversity hotspot, is particularly poorly studied. The aim of this paper is to produce a first annotated checklist of the wild bees of Lebanon from new and museum collections, literature records, and verified occurrences from online databases. The present list totals 573 species for Lebanon of which 289 are reported for the first time, but the estimated diversity is likely to be closer to 700. Preliminary information on local distributions and flower records are also presented. The local species assemblages indicate affinities with montane habitats of the Mediterranean and Anatolia and the semi-arid habitats of the Levant and north Africa. This study also encourages further research on local wild bee faunas and the use of this knowledge for conservation purposes.

A survey on Megachilidae (Hymenoptera, Apoidea) species available in Iranian Pollinator Insects Museum of Yasouj University

Article

Full-text available

Mar 2021

In this study, the distribution of 88 species of the family Megachilidae, out of 3678 specimens in the Iranian Pollinator Insects Museum of Yasouj University (IPIM), which were collected in spring and summer 2009–2017 from different regions of Iran including Ardabil, Chaharmahal-o Bakhtiari, Fars, Golestan, Isfahan, Kerman, Khuzestan, Kohgiluyeh-va Boyer-Ahmad, Sistan-o Baluchestan, Alborz, are presented. Data on the number of specimens, locations, coordinates and distribution maps for Iran and global distribution of all species (where available) are also provided. Megachile (Megachile) octosignata Nylander 1852 is first recorded from Iran. Ninetheen species are reported for the first time from Chaharmahal-o Bakhtiari and Kohgiluyeh-va Boyer-Ahmad Provinces.

Sur la présence en France d’Andrena confinis Stöckert, 1930, et d’Osmia bidentata Morawitz, 1876 (Hymenoptera, Andrenidae, Megachilidae)

Article

Dec 2020

Romain Le Divelec

On the presence in France of Andrena confinis Stöckert, 1930, and Osmia bidentata Morawitz, 1876 (Hymenoptera, Andrenidae, Megachilidae). Andrena confinis Stöckhert, 1930, and Osmia bidentata Morawitz, 1876, are two wild bee species for which presence in France is dubious. Some French specimens were found in the collections of the Muséum national d’Histoire naturelle (Paris), so affiliation of these two species to the French fauna is confirmed. Major diagnostic features, ecology and distribution are provided.

Field Guide to the Bees of Great Britain and Ireland (Bloomsbury)

Cover Page

Full-text available

Nov 2015

Steven Falk

A comprehensive guide to the bees of Great Britain, Ireland and the Channel Islands covering all 275 species on the British list in 2015. The book can be purchased from most of the larger natural history book suppliers. 432 pp. Bloomsbury British Wildlife Field Guides. ISBN: 978-1-910389-02-7 (hardback), ISBN: 978-1-910389-03-4 (paperback).

Catalogo degli Imenotteri italiani. IV (Apoidea: Colletidae, Andrenidae, Megachilidae, Anthophoridae, Apidae).

Article

Full-text available

Jul 1994

Guido Pagliano

Check-list of Hymenoptera in Italy. IV. Apoidea: Colletidae. Andrenidae, Megachilidae, Anthophoridae and Apidae. A list of the species of Italian apoidea has been made. Data from bibliography and from numerous Italian collections have been deduced. An index in alphabetical order of species, subspecies and synonymies follows. 43 taxa are new for Italy

Ergänzungen zur Bienenfauna (Hymenoptera: Apidae) von Wien, Österreich

Article

Full-text available

Jan 2016

Additions to the bee fauna (Hymeno ptera: Apidae) of Vienna, Austria. – As an addition to a recently published checklist of the bees in Vienna, six first records are communicated: Hylaeus pectoralis Förster, 1871, Colletes mlokossewiczi radsZoWski, 1891, Andrena dorsalis brullé, 1832, Anthidium florentinum (Fabricius, 1775), Megachile flabellipes PéreZ, 1895, and Hoplos-mia ligurica (moraWitZ, 1868). Three species with previously doubtful records are confirmed to occur in Vienna: Dioxys tridentata (nylander, 1848), Hoplosmia bidentata (moraWitZ, 1876), and Tetralonia salicariae (lePeletier, 1841). This increases the number of species recorded from Vienna from 456 to 465. The presence of Anthidium florentinum in Austria is herewith confirmed. Further new records of the following regionally rare bee species are reported: Lasioglossum pallens (brullé, 1832), Sphecodes majalis PéreZ, 1903, Nomioides minutissimus (rossi, 1790), Hoplitis claviventris (thomson, 1872), Hoplitis papaveris (latreille, 1799), Lithurgus cornutus fuscipennis lePeletier, 1841, and Ceratina acuta Friese, 1896. In addition Colletes mlokossewiczi is recorded from Lower Austria for the first time. Zusammenfassung In Ergänzung zur kürzlich erschienenen Checkliste der Bienen Wiens werden sechs Erstnach-weise mitgeteilt: Hylaeus pectoralis Förster, 1871, Colletes mlokossewiczi radsZoWski, 1891, Andrena dorsalis brullé, 1832, Anthidium florentinum (Fabricius, 1775), Megachile flabellipes PéreZ, 1895 und Hoplosmia ligurica (moraWitZ, 1868). Drei Arten, die bisher aufgrund unklarer Fundortangaben für Wien fraglich waren, können nun bestätigt werden: Dioxys tridentata (ny-lander, 1848), Hoplosmia bidentata (moraWitZ, 1876) und Tetralonia salicariae (lePeletier, 1841). Dies erhöht die Zahl der sicher aus Wien nachgewiesenen Bienenarten von 456 auf 465. Das Vorkommen von Anthidium florentinum in Österreich ist somit erstmals belegt. Weiters wird über interessante Wiederfunde der folgenden regional seltenen Bienenarten berichtet: Lasioglossum

Specialist bees collect Asteraceae pollen by distinctive abdominal drumming (Osmia) or tapping (Melissodes, Svastra)

Article

Full-text available

Jun 2017
ARTHROPOD-PLANT INTE

Jim Cane

Four species of western US Osmia (3 Cephalosmia) that are Asteraceae specialists (mesoleges) were observed using a stereotypical means of collecting pollen—abdominal drumming—to gather pollen from 21 flowering species representing nine tribes of Asteraceae. Abdominal drumming is a rapid dorso-ventral motion of the female’s abdomen (467 pats/min) used to directly collect and place pollen in the bee’s ventral scopa. A co-occurring generalist, O. lignaria, never drummed Asteraceae flowers for pollen, but instead used its legs to harvest pollen. Observed drumming by several other osmiines is noted. A different pollen-harvesting behavior, abdominal tapping, is described for two eucerine bees (Melissodes agilis and Svastra obliqua), both oligolectic for the Asteraceae. The behavior also involves a dorso-ventral motion, but they tap their distal abdominal venter against disk flowers at a slower tempo (304 taps/min). These females’ distal sternites have distinctly dense and long hair brushes for acquiring pollen by this behavior. Brief accounts of similar abdominal pollen gathering behaviors by other megachilids are summarized.

Pollen preferences of Hoplosmia bidentata and Lithurgus cornutus (Hymenoptera: Megachilidae)

Article

Full-text available

Sep 2007

In this study pollen from the scopae of two species of solitary bees was identified from five sites in Turkey in the vicinity of the Central Anatolian cities Ankara, Eskişehir and Çankiri. The samples of Lithurgus cornutus (Fabricius) and Hoplosmia bidentata (Morawitz) (Hymenoptera: Megachilidae) were collected between June and August. Most of the 162 specimens contained more than one type of pollen. L. cornutus (Fabricius) was provisioned with pollen from eight families, H. bidentata (Morawitz) from eleven families. New types of pollen for both species were also identified.

Weather and the use of empty gastropod shells by arthropods

Article

Full-text available

Jul 2008

Arthropods frequently use empty snail shells as shelter or nesting sites. This study analyses the use of shells from the arid-dwelling land snail Sphincterochila candidissima (Draparnaud 1801) by arthropods in the Sierra Elvira (south-eastern Spain). Arthropods of 11 different orders occupied 15.6% of shells. Salticidae (54.6%) and Apoidea (21.3%) comprised the majority of arthropods. Shells were used primarily when environmental temperatures were lowest, suggesting that, in the Sierra Elvira, arthropods use shells to escape from the cold. Weather explained 73% of the variation in shell occupation. However, Apoidea used shells independently of weather, occupying shells as nesting chambers. These results suggest that snail shells may be important refugia and nesting sites for arthropod fauna.

Die Leucospidae (Hymenoptera: Chalcidoidea) der Schweiz, mit einem Bestimmungsschlüssel und Daten zu den europäischen Arten

Article

Full-text available

Jan 2000
REV SUISSE ZOOL

The Leucospidae (Hymenoptera: Chalcidoidea) of Switzerland, with a key and data on the European species. - In this paper the taxonomy and distribution of the Swiss species of Leucospis Fabricius are presented. Four of the seven European species are recorded for Switzerland: L. biguetina Jurine, L. dorsigera Fabricius, L. gigas Fabricius, and L. intermedia llliger. Except for L. dorsigera the main distribution lies in the Southern parts of the country. The altitudinal gradient ranges from the colline to the subalpine belt. Adults appear from May to September. L. biguetina and L. gigas are considered vulnerable with regard to the Swiss fauna. Almost exclusively females of L. biguetina and L. gigas are known from the study area, hence these species are likely to reproduce by thelytoky. A key, diagnoses, morphometric and distributional data are also given for the other two European species, L. bifasciata Klug and L. brevicauda Fabricius. The hosts of all species are critically reviewed and new hosts are recorded for L. dorsigera and L. gigas.

Les Osmies de la Faune Française: [Hymenopt. Apidae]

Article

Aug 2023

Raymond Benoist

Notes détachées sur l'instinct des Hyménoptères mellifères et ravisseurs avec la description de quelques espèces

Article

C. Ferton

Checklist of Hungarian Sphecidae and Apidae species (Hymenoptera, Sphecidae and Apidae)

Article

Jan 2011

Zsolt Józan

H-7453 Mernye Rákóczi út 5., Hungary JóZan, Zs.: Checklist of Hungarian Sphecidae and Apidae species (Hymenoptera, Sphecidae and Apidae). Abstract: Checklist of Hungarian sphecoid wasps (Sphecidae) and bees (Apidae). 306 Sphecidae and 704 Apidae species are listed. Species, which were collected 50 years or earlier, treated as extinct species. I have not seen voucher specimen of the following species: Halictus varipes, Halictus carinthiacus, Hylaeus hungari-cus, Osmia hybrida, Osmia versicolor, Sphecodes hungaricus, Sphecodes schencki and Sphecodes sareptensis. These species are not listed. Belomicroides fergusoni (Sphecidae), Andrena pyropygia, Halictus cochlearitar-sis, Melitta wankowiczi and Osmia scutellaris are first records for Hungary.

Palaearctic Osmia bees of the subgenus Hoplosmia (Megachilidae, Osmiini): Biology, taxonomy and key to species

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