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Belgian Journal of Entomology 105: 124 (2020) ISSN: 2295-0214
Belgian Journal of Entomology
Lasioglossum dorchini (Hymenoptera: Apoidea: Halictidae)
a new species of bee from Israel
Alain PAULY1, Karmit LEVY2, Grégoire NOËL3, Gontran SONET1 , Jean-Luc BOEVÉ1
1 Royal Belgian Institute of natural Sciences, O.D. Taxonomy & Phylogeny, Rue Vautier 29, 1000 Brussels,
Belgium; e-mails:, (corresponding author)
2 The Hebrew University of Jerusalem, Faculty of Agriculture, Food and Environment, Rehovot, Israel
3 Functional and Evolutionary Entomology, Université de Liège - Gembloux Agro-Bio Tech, Passage des
déportés 2, B-5030 Gembloux, Belgium.
Published: Brussels, 27 November 2020
PAULY et al. A new species of bee from Israel
Citation: PAULY A., LEVY K., NOËL G., SONET G., BOEVÉ J.-L. & MANDELIK Y., 2020. - Lasioglossum dorchini
(Hymenoptera: Apoidea: Halictidae), a new species of bee from Israel. Belgian Journal of Entomology, 105: 1
ISSN: 1374-5514 (Print Edition)
ISSN: 2295-0214 (Online Edition)
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Front cover: Head of a female of Lasioglossum dorchini sp. nov., paratype, Israel: Nahal Alexander. © A. Pauly.
Belgian Journal of Entomology 105: 124 (2020)
Lasioglossum dorchini (Hymenoptera: Apoidea: Halictidae)
a new species of bee from Israel
Alain PAULY1, Karmit LEVY2, Grégoire NOËL3, Gontran SONET1, Jean-Luc BOEVÉ1
1 Royal Belgian Institute of natural Sciences, O.D. Taxonomy & Phylogeny, Rue Vautier 29, 1000 Brussels,
Belgium; e-mails:, (corresponding author)
2 The Hebrew University of Jerusalem, Faculty of Agriculture, Food and Environment, Rehovot, Israel
3 Functional and Evolutionary Entomology, Université de Liège - Gembloux Agro-Bio Tech, Passage des
déportés 2, B-5030 Gembloux, Belgium.
This paper describes a new species, Lasioglossum dorchini, occuring in sand dunes in Israel. It
is close to Lasioglossum leptocephalum. Its phylogenetic relationships with the other species
of the virens/littorale group are analyzed.
Keywords: bees, Israel, Lasioglossum, new species, sand dunes
Cette publication donne la description d'une nouvelle espèce, Lasioglossum dorchini,
découverte dans les dunes sableuses en Israël. Elle est proche de Lasioglossum leptocephalum.
Ses relations phylogénétiques avec les autres espèces du groupe virens/littorale sont analysées.
Israel and Palestine alone are home to approximately 1100 known bee species (O'TOOLE &
RAW, 1991). Extensive collecting throughout Israel in recent years has led to the discovery of
new species of wild bees (PISANTY et al., 2016). In this paper we describe an additional new
species from Israel, Lasioglossum dorchini Pauly sp. nov., wich is very close and sympatric to
the known species Lasioglossum leptocephalum (Blüthgen, 1923). The new species has been
recently discovered in bee surveys in Nahal Alexander National Park in the central coastal Plain
of Israel where it is the most common Halictid species collected in pan traps (i.e. colored bowls
containing soapy water).
The genus Lasioglossum has a complex and unstable subgeneric classification and, although
recent phylogenetic and molecular studies have been published (GIBBS et al., 2013; GIBBS,
2018), these do not yet fully cover groups of Palaearctic species. It was therefore premature to
adopt this new classification here because it would have forced us to make many extrapolations
for species of ambiguous position and not yet sequenced. BYTISNKY-SALZ & EBMER (1974)
placed L. leptocephalum in the paraphyletic subgenus "Evylaeus Robertson, 1902". The species
belongs to the Halictus virens species-group distinguished by BLÜTHGEN (1931: 393) and to
the Lasioglossum littorale group of EBMER (1974: 140; 1982: 218) (= L. virens group of EBMER
1976: 244; 1993: 780). All these species are classified by PESENKO (2007: 26) in his new
subgenus "Virenshalictus", including ten species. Preliminary molecular studies suggest that
this group of species should be included in the subgenus Hemihalictus Cockerell, 1897 as
delimited by GIBBS et al. (2013) and not in the subgenus Dialictus Robertson, 1902 that is
largely comprised of species with metallic green reflections but with carinate propodeum.
PAULY et al. A new species of bee from Israel
Here, we aim to (i) understand if L. dorchini is one a well-delineated species on the basis of
morphological and molecular analyses; and (ii) consider the phylogenetic relationships among
different known species of the virens/littorale group of species.
Material and methods
Revision of types and identification of non-type specimens was based on material from the
following institutions (with their acronyms): The Steinhardt Museum of Natural History, Tel
Aviv University (SMNHTAU); Museum fr Naturkunde an der Humboldt Universität zu
Berlin, Germany (MNHUB); Natural History Museum, London, UK (NHML); Royal Belgian
Institute of Natural Sciences, Brussels, Belgium (RBINS). A total of 475 specimens were
examined. In addition, a DNA fragment of the cytochrome c oxidase subunit I (COI) gene was
sequenced for recently collected specimens of the following species of the virens/littorale
group: Lasioglossum aureolum (Pérez, 1903), L. dorchini Pauly sp. nov., L. leptocephalum
(Blüthgen, 1923), L. littorale (Blüthgen, 1923) and L. aff littorale, L. musculoides (Ebmer,
1974), L. pseudoleptocephalum (Blüthgen, 1923), L. pseudolittorale (Blüthgen, 1923),
L. virens (Erichson, 1835) (see acknowledgments for the collectors).
For molecular analyses, we followed the protocol from PAULY et al. (2015) and PAULY et al.
(2019) for specimen samples AP221, AP303, AP304, AP699, AP700, AP701, AP702, AP703,
AP704 and AP705 (Table 1). Briefly, one middle leg per specimen was used to extract DNA
with the NucleoSpin® Tissue Kit (Macherey-Nagel, Germany) following instructions of the
manufacturer. One pair of primers (LCO1490 and HCO2198) was used to amplify the
cytochrome c oxidase subunit I (COI) mitochondrial gene, referred to as the DNA barcode for
animals (FOLMER et al., 1994). The contigs resulting from the amplification of both primers
were aligned and edited in CodonCode Aligner© ver. 5.0.1 (CodonCode Corp., Centerville,
Massachusetts). In addition, barcode sequences of specimen samples AP404, AP409, AP410,
AP454, AP463, AP499, AP500, AP501, AP502 and AP503 (Table 1), were produced and
analysed at the Canadian Centre for DNA Barcoding (CCBD) using the protocols described in
IVANOVA et al. (2006) and DEWAARD et al. (2008). The resulting consensus sequences (n = 20
species) with range of 482 676 bp were subject to BLAST search to check for possible
contaminations. To complete the dataset, we added the barcodes KJ839379 and KJ836462 of
L. virens from the work of SCHMIDT et al. (2015).
We selected four unique barcode sequences, KJ839608 and KJ839285 for Lasioglossum
nitidulum (Fabricius 1804), and KJ839826 and GU706057 for Lasioglossum morio (Fabricius,
1793) (SCHMIDT et al., 2015), retrieved from GenBank to constitute the outgroup rooting the
generated phylogenetic trees. These two species belong to the subgenus Dialictus, a group with
metallic reflections close to our species. We aligned all the DNA sequences using ClustalW
(LARKIN et al., 2007) with the default parameters in MEGA ver. 7.0.21 (KUMAR et al., 2016).
On the basis of the global similarity of the nucleotide sequences, we reconstructed a Neighbour-
Joining (NJ) tree using MEGA ver. 7.0.21 with bootstrap pseudo-replicates (n = 1000).
For phylogenetic reconstruction based on bayesian inference (BI) method, we used
PartitionFinder2 ver2.1.1 (LANFEAR et al., 2017) to explore best nucleotide substitution model
for our aligned DNA dataset by their nucleotide positions. The TIM gamma model of rate
heterogeneity (TIM + G), the TrN (TRN + I) and the F81 (F81 + I) with invariables sites
substitution models were selected as best nucleotide substitution models for nucleotide position
Belgian Journal of Entomology 105: 124 (2020)
1, 2 and 3 respectively. We set-up Markov chain Monte Carlo (MCMC) length to five million
generations in two parallel runs. After the run, we checked the posterior probabilities as well as
the effective sample size (> 300) with Tracer ver. 1.7.1 program (RAMBAUT et al., 2018).
Finally, we summarized BI into phylogenetic tree via TreeAnnotator ver. 2.5.2. (included in
BEAST environment) program with a burn-in of 25%. All phylogenetic trees were compiled
using FigTree ver. 1.4.3 (RAMBAUT, 2017) and the document (.svg) was illustrated with
Inkscape ver. 0.92.2.
Fig. 1. Lasioglossum dorchini, Israel: Nahal Alexander. a, female; b, male.
Lasioglossum dorchini Pauly sp. nov.
DIAGNOSIS. Close to L. leptocephalum. The female differs by the less slender head (Fig. 2), the
larger clypeus, the first tergum more densely punctuated in the middle (sparsely punctated in
the middle in L. leptocephalum) (Fig. 3), the third tergum pubescent only on its base, with very
fine setae on the apical half (nearly completely covered with plumose setae in
L. leptocephalum) (Figs 4, 5), the legs dark brown (pale brown in L. leptocephalum) (Fig. 6),
the less long propodeum (Fig. 7d versus Fig. 11d). The male differs from L. leptocephalum
(which is known only by the type series of L. krugeri Blüthgen, 1930) by the less abundant
tomentum and the shorter erect setae on the terga (Fig. 8e compared to Fig. 12d) and the shorter
propodeum (Fig. 8c compared to Fig. 12c).
Colouration. Body with metallic bronze-green reflections, apical margins of terga largely
amber (Figs 1a, 4b), sometimes almost entirely amber; sterna completely amber (Fig. 6b). Legs
black, end of the femurs and basal half of the basitarsi of the hind legs brown (Fig. 6b).
Flagellum brown below. Mandibles dark brown. Tegulae pale yellow translucent (Fig. 7b, c).
Structure, punctation and pubescence. Head very elongate (length/width = 1.19) (Figs 2b, 7a).
Clypeus and supraclypeal area long (Fig. 2b); vertex narrow (vertex width/ intertegular distance
= 0.68), finely striated (Fig. 7b); frons densely and finely punctuated (Fig. 2b); apical half of
the clypeus and lower adjacent part of the eyes black and smooth, with sparse punctures (Fig.
2b); genae densely striate. Mesosoma. Pronotum angles obtuse. Scutum and scutellum densely
and finely punctuated, puncture interspaces equal a point diameter, the surface dull and
completely striated (Fig. 7c). Propodeum not carinate, short (propodeal/metanotal length = 1.4),
wrinkled on the basal half, micro-tessellated on the apical half (Fig. 7d). Metasoma. First
PAULY et al. A new species of bee from Israel
Table 1. Female specimen information from the virens / littorale group with their GenBank accession or BOLD
process ID.
Lasioglossum aureolum
D. Genoud
Israel, Nahal Alexander
K. Levy
Lasioglossum dorchini
Israel, Nahal Alexander
K. Levy
Lasioglossum aureolum
Portugal, Tavira, Cabanas
A. Livory & R.
Lasioglossum aff littorale
Portugal, Barra
A. Livory & R.
Lasioglossum littorale
Italy, Toscana, Parco regional
(Migliarino san Rossore
M. Boschetti
Portugal, Algarve, Praia do Barril,
T. Wood
Lasioglossum aff littorale
Italy, Sardarnia, Cagliari,
Castiadas, San Giusta
E. Dufrêne
Israel, Park Britannia
T. Chaprazaro
Lasioglossum dorchini
Israel, Nahal Alexander
K. Levy
Lasioglossum dorchini
Israel, Nahal Alexander
K. Levy
Lasioglossum dorchini
Israel, Nahal Alexander
K. Levy
Lasioglossum dorchini
Israel, Nahal Alexander
K. Levy
Israel, Holot Shunera
G. Pisanty
Israel, Holot Shunera
G. Pisanty
Lasioglossum musculoides
Morocco, Haddada
I. El Abdouni
Lasioglossum aff littorale
Morocco, Bouknadel
I. El Abdouni & P.
Lasioglossum aff littorale
Morocco, Haddada
I. El Abdouni & P.
Lasioglossum aff littorale
Morocco, Bouknadel
I. El Abdouni & L.
Lasioglossum aff littorale
Morocco, Haddada
I. El Abdouni & A.
tergum finely and rather densely punctuated, puncture interspaces equal two puncture
diameters, the inclined base finely striate (Fig. 3b). Sides of inclined base of tergum 1 and on
the bases of terga 2 to 4 with grey plumose tomentum (Fig. 4b). Middle of the tergum 3 with
short weakly branched setae, without plumose hairs (Fig. 5b). Legs. Hind tibial spur with 3 long
oblique teeth (Fig. 7e).
MALE. Length 6 mm.
Colouration. Head and mesosoma with blue-green metallic reflections, metasoma black
without or with very weak metallic reflections (Fig. 1b). Apical margins of terga hyaline fulvous
(Fig. 8e). All tarsi, basal and apical parts of hind tibiae pale yellow (Fig. 8f). Apical third of the
clypeus pale yellow (Fig. 8a). Underside of flagellum ocraceous (Fig. 8g). Tegulae pale yellow
translucent (Fig. 8b).
Structure, punctation and pubescence. Head long (length/width=1.12). Flagellomeres 1,5 time
longer than wide (Fig. 8g). Mesosoma. Scutum and scutellum densely and finely punctuate,
puncture interspaces dull and equal to a puncture diameter (Fig. 8b). Propodeum not carinate,
the propodeal area relatively short (propodeal/metanotal length=1.25), with some wrinkles
reaching posterior margin (Fig. 8c). Metasoma. Surface of the terga shiny, with punctures fine
and comparatively dense (puncture interspaces equal 1.52 puncture diameter) (Fig. 8d, e).
Belgian Journal of Entomology 105: 124 (2020)
Fig. 2. Comparison of the heads; a, Lasioglossum leptocephalum, Israel: Araba; b, Lasioglossum dorchini, Israel:
Nahal Alexander.
PAULY et al. A new species of bee from Israel
Fig. 3. Comparison of the punctuation in the middle of the first tergum; a, Lasioglossum leptocephalum, Israel:
Park Britania; b, Lasioglossum dorchini, female, Israel: Nahal Alexander.
Belgian Journal of Entomology 105: 124 (2020)
Fig. 4. Comparison of the pilosity on the metasoma; a, Lasioglossum leptocephalum, Israel: Park Britannia;
b, Lasioglossum dorchini, female, Israel: Nahal Alexander.
PAULY et al. A new species of bee from Israel
Fig. 5. Comparison of the pilosity on the third tergum; a, Lasioglossum leptocephalum, Israel: Park Britannia;
b, Lasioglossum dorchini, female, Israel: Nahal Alexander).
Belgian Journal of Entomology 105: 124 (2020)
Fig. 6. Comparison of the colour of the hind leg; a, Lasioglossum leptocephalum, Israel: BatYam; b, Lasioglossum
dorchini, Israel: Nahal Alexander.
PAULY et al. A new species of bee from Israel
Fig. 7. Lasioglossum dorchini, Nahal Alexander; a, head; b, vertex; c, scutum and scutellum; d, propodeum;
e, calcar.
Sterna with oblique setae apically (Fig. 8f). Genitalia. Gonocoxites with gonostyli and ventral
membranes illustrated in Fig. 8h.
DISTRIBUTION AND ECOLOGY. Lasioglossum dorchini is known only from the central coastal
plain of Israel (one specimen from Acre in the northern coastal plain). The type locality in Nahal
Alexander National Park is characterized by semi-stabilized sand dunes typical of Israel’s
central coastal plain (KUTIEL, 2001) (Figs 1518). The park was planted in part with non-native
Eucalyptus trees in the mid 1960s, mainly E. camaldulensis Dehnh. The natural shrubland flora
is dominated by Artemisia monosperma Delile, Retama raetam (Forssk.) Webb and Ephedra
aphylla Forssk. Bees were collected in the north western part of the park, during spring
(February-April). Most specimens were captured with pan traps and some others were collected
with nets while visiting flowers of Asteraceae (Crepis aculeata (DC.) Boiss., Senecio glaucus
L., and S. leucanthemifolius subsp. vernalis (Waldst. & Kit.) Greuter).
Belgian Journal of Entomology 105: 124 (2020)
Fig. 8. Lasioglossum dorchini, male, Nahal Alexander; a, head; b, scutum and scutellum; c, propodeum; d, first
tergum; e, metasoma; f, hind leg and pilosity of the sterna; g, antenna; h, genitalia.
PAULY et al. A new species of bee from Israel
TYPE MATERIAL (456 specimens; SMNHTAU, RBINS). Holotype : ISRAEL, Nahal
Alexander, 32.401 34.882, 1.V.2016, leg. K. Levy (AP 304) (GenBank MT221620)
Paratypes. ISRAEL, Nahal Alexander, 31.III.2016, 1, 1; 3.IV.2016, 1; 5.IV.2016, 3,
27; 6.IV.2016, 1, 9; 7.IV.2016, 4; 20.IV.2016, 1; 21.IV.2016, 1, 22; 24.IV.2016,
6, 30; 25.IV.2016, 10; 1.V.2016, 1; 5.V.2016, 1; 4.III.2017, 5; 9.III.2017, 4;
10.III.2017, 35; 23.III.2017, 2; 31.III.2017, 1; 3.IV.2017, 4; 4.IV.2017, 23; 6.IV.2017,
10; 10.II.2018, 14; 11.II.2018, 46; 14.II.2018, 30; 15.II.2018, 12; 16.II.2018, 87;
7.III.2018, 10; 8.III.2018, 7; 9.III.2018, 1; 16.III.2018, 1; 17.III.2018, 2; 4.IV.2018,
1, 5; 5.IV.2018, 2; 6.IV.2018, 2, 14; 7.IV.2018, 1, 4; 8.IV.2018, 1, all. leg. K.
Levy (SMNHTAU, RBINS). Giv'at Homera Nature Reserve, 31.935 34.743, 6.II.2015, pan
trap, 2, leg. G. Pisanty (202534, 202559) (SMNHTAU). Palmahim, 31.927 34.730,
6.II.2015, pan trap, 2, leg. G. Pisanty (202520) (SMNHTAU). 'En Sarid, 32.273 34.928,
5.IV.2012, leg. O. Afik (SMNHTAU). Cesarea, 30.IV.1963, 1, leg. Kugler (SMNHTAU).
Holon, 1.III.1986, 1, leg. E. Shney-Dor (SMNHTAU). Holon, 32.002 34.787, 4.V.1978,
1, leg. M. Kaplan (SMNHTAU). Holon Dunes, 31.998 34.787, 1.III.1986, 1, leg. E.
Shney-Dor (SMNHTAU). Tel-Aviv Swamp, 31.935 34.743, 9.IV.1981, 1, leg. Freidberg
(SMNHTAU). Acre, 30.IV.1963, 1, leg. Kugler (SMNHTAU). Berekhat Ya'ar, North,
32.412 34.898, 23.V.2003, 1, leg. A. Freidberg (SMNHTAU). Nizzanim, 31.722 34.603,
13.V.2003, 1, leg. L. Freidman (SMNHTAU). 17 km SSW Tel Aviv, Dünen W Kefar,
N31.41 E35.28, 13.V.1996, 1, leg. O. Niehuis (col. Ebmer) (specimen examined and
identified by Ebmer as L. sinaiticum). Sharon Plain, Netanya shore, 32.286 34.841, 1,
27.IV.2009, at 9 am, leg. A. Dorchin.
ETYMOLOGY. This new species is dedicated to Achick Dorchin, bee specialist at the Tel Aviv
University, who discovered the first specimen I examined of this species.
Lasioglossum leptocephalum (Blüthgen, 1923)
Halictus leptocephalus BLÜTHGEN, 1923: 245. Lectotype : Tunisia, Nefta, 14.V.1913 (coll.
Blüthgen, MNHUB).
= Lasioglossum sinaiticum BYTINSKI-SALZ & EBMER, 1974: 195. Holotype : Sinai, Nahal
Yam (lagoon of Bardawil), 1.II.1973, leg. A. Freidberg (coll. Ebmer, in Linz) (not examined).
Paratypes: 3, idem (Linz and col. Bytinsky-Salz) (2 examined in SMNHTAU), syn. nov.
= ?Halictus krugeri BLÜTHGEN, 1930: 222. Holotype : Libya, Agedabia (SW Tripoli),
20.V.1925, leg. Krüger (MNHUB) (examined). Paratypes: 4, idem (MNHUB and R.U.
Agrario Bengasi) (examined in MNHUB).
NOTE. WARNCKE (1982: 69) erroneously considered L. leptocephalum as a subspecies of
L. albovirens (Pérez, 1895) with a more elongated head and L. sinaiticum as a simple form of
L. albovirens leptocephalum. He also placed L. krügeri in synonymy with L. albovirens
leptocephalum. The synonymy of L. krugeri remains doubtful and should be confirmed by the
discovery of the two sexes in a same locality; it is mentioned here with a "?".
DISTRIBUTION AND ECOLOGY. Lasioglossum leptocephalum occurs in coastal dunes from
Tunisia to Israel in the east. The locality in Holot Shunera is a partially stabilized sand dune
ecosystem in the Western Negev (Gideon Pisanty, personal communication). In Libya, it has
been collected in stabilised sands of the green belt.
Belgian Journal of Entomology 105: 124 (2020)
Fig. 9. Lasioglossum leptocephalum; a, b, female lectotype (Nefta); c, e, Djerba; d, Tripoli; f, Bat Yam.
PAULY et al. A new species of bee from Israel
Fig. 10. Lasioglossum sinaiticum, female paratype, Nahal Yam, (syn. of L. leptocephalum); a, head; b, scutum;
c, propodeum; d, first tergum; e, metasoma; f, hind leg; g, labels.
Belgian Journal of Entomology 105: 124 (2020)
Fig. 11. Lasioglossum krugeri, male holotype; a, head; b, scutum and scutellum; c, propodeum; d, metasoma;
e, antenna.
MATERIAL (19 specimens). TUNISIA. Tozeur, 10.V.1913, 1 (paratype) (col. Blüthgen,
MNHUB) (BLÜTHGEN, 1923: 245; 1924: 307). Tunis, "Ued Mda" (Oued Melah), 1, col.
Alfken (? MNHUB) (BLÜTHGEN, 1923: 245; 1924: 307). Nefta, 14.V.1913, 2 (BLÜTHGEN,
1924: 307). Tunis, 1 (MNHUB). Gabès, 1 (MNHUB) (BLÜTHGEN, 1934: 190).
LIBYA. Tripolitania, 9 km S. of Tripoli, 18.III.1951, fixed sandy ground in green belt, 1, leg.
K.M. Guichard (NHML).
EGYPT. Mansuria, 30.XII.1926, 1 (BLÜTHGEN, 1934: 190). Araba, N. Sinai, 3.II.1973, 3,
leg. D. Furth (= Wadi Araba) (29.574779, 34.979854) (SMNHTAU).
ISRAEL. Park Britannia, 31.707 34.921, 17.IV.2016, 1, leg. T. Chaprazaro (251704)
(SMNHTAU). Bat Yam, 32.008 34.898, 2.III.1919, 6, leg. Bytinsky Salz (MNHUB).
Holot Shunera, 30.942 34.597, 25.II.2015, 2, in stabilized sand dune, leg. G. Pisanty (205506)
PAULY et al. A new species of bee from Israel
Fig. 12. Distribution map of Lasioglossum leptocephalum.
Table 2. Morphological comparison between Lasioglossum leptocephalum and L. dorchini
Characters and measurements
Lasioglossum leptocephalum
Lasioglossum dorchini
HL/HW= head length/ width
1.23 (Fig. 2a), narrower
1.14 (Fig. 2b), wider
CL/CW= clypeus length/ width
0.67 (Fig. 2a), narrower
0.61 (Fig. 2b), wider
UID/LID= upper interorbital distance/
lower interorbital distance
1.14, more convergent below
(Fig. 2a)
1.11, less convergent below
(Fig. 2a)
punctuation of middle of first tergum
sparse (Fig. 3a)
dense (Fig. 3b)
pubescence of third tergum
plumose setae covering 2/3 of
the basal surface
(Fig. 5a)
plumose setae delimited to the
sides, short and unbranched in
the middle (Fig. 5b)
length of the propodeal area/ length of
the metanotum
1.5 (Fig. 10c)
1.4; (Fig. 7d)
colour of the hind leg
with more extended pale
brown maculations (Fig. 6a)
nearly completely dark brown
(Fig. 6b)
(type of Halictus krugeri
Blüthgen 1930)
(males from Nahal
setae of the terga
long, white and plumose
(Fig. 11d)
short, grey and weakly
branched (Fig. 8e)
colour of the metasoma
with metallic blue reflections
(Fig. 11d)
mostly black (Fig. 8e)
length of the propodeal area / length of
the metanotum
1.66 (Fig. 11c)
1.25 (Fig. 8c)
wrinkles of the propodeal area
wrinkles limited to basal half
of the propodeal area (Fig.
wrinkles reaching posterior
margin of propodeal area
(Fig. 8c).
Belgian Journal of Entomology 105: 124 (2020)
Phylogenetic analysis
All specimens identified as L. dorchini based on morphological examination formed a
monophyletic group with maximal posterior probability and bootstrap support values in all BI,
NJ and ML phylogenetic analyses, respectively (Figs 13, 14). Similarly, specimens of L. virens,
L. leptocephalum and L. aureolum were all reciprocally monophyletic (Figs 13, 14). Our results
further show that specimens identified as L. littorale or aff. littorale (AP409, AP410, AP463,
AP702, AP703, AP704 and AP705) were polyphyletic, comprising at least four different
lineages across the virens/littorale group phylogeny (Figs 13, 14). Phylogenetic relationships
among the different species of the virens/littorale group remain unsupported by our phylogeny
(Figs 13, 14).
Fig. 13. Neighbour-joining tree reconstructed using 22 barcode sequences (658bp) of the cytochrome oxidase c
subunit I gene for specimens currently identified into the virens/littorale group of species. The tree is drawn to
scale, with branch lengths representing p-distances (i.e. proportions of variable sites). Four barcodes sequences
(shaded in grey) are used as outgroup for the phylogenetic tree: Lasioglossum morio and L. nitidulum. Each label
corresponds to the study code (Table 1) or GenBank accession followed by the species name and the country of
collection. The shaded colours on the tree correspond to morphological delineation except for the species identified
as Lasioglossum littorale or aff. littorale which are left unshaded. Values at node correspond to bootstrap values
Discussion and conclusions
Lasioglossum dorchini is a species so close to L. leptocephalum that it may be considered as a
simple variation. Here however, integrative taxonomy combining barcodes and morphological
features support its validity as a new species.
PAULY et al. A new species of bee from Israel
Fig. 14. Phylogenetic tree constructed using Bayesian inference. The tree is based on 22 barcode sequences
(676 bp) of the cytochrome oxidase c subunit I gene for specimens currently identified into the virens/littorale
group of species. Four barcodes sequences (shaded in grey) are used as root for the phylogenetic tree: Lasioglossum
morio and Lasioglossum nitidulum. Each label corresponds to the study code (Table 1) or GenBank accession
followed by the species name and the country of collection. Posterior probabilities are given at nodes. The shaded
colors on the tree correspond to morphological delineation except for the species identified as Lasioglossum aff
littorale which was let as blank. Only posterior probabilities above 0.95 are showed in the figure.
Lasioglossum littorale has been considered as a widespread species occuring in coastal habitats
of the Mediterranean Bassin. It has been split by EBMER (1972, 1976) in different insular and
geographic subspecies that are difficult to identify. The morphological heterogeneity of the
different subspecies with respect to body size or length of the head complicates their
classification. Our molecular analyses demonstrate that the species is composite. Additional
freshly collected material, especially from Southern Spain and North Africa, is needed to clarify
the systematics of the all the species provisonally identified as "aff littorale". But our
preliminary revision of the barcoded specimens already reveals some subtle morphological
differences, for instance, in the density and size of the punctation of the scutum and terga.
All species of the virens/littorale group occur in sandy habitats, and particulary in coastal dunes,
a type of habitat that is strongly threatened by infrastructures development for tourism and
urbanisation. Consequently, the geographic distribution of these species is very limited, despite
the fact that these specialized species are often counted among the most abundant bees collected
with pan traps in these particular habitats (BOSCHETTI et al., 2017).
The subgeneric classification of the Lasioglossum Hemihalictus series remains challenging, and
is currently been revised based on molecular phylogenetic evidence. It would be premature to
recognise the virens/littorale group as a valid subgenus before this group is proved
monophyletic. To achive this, additional non-metallic species (that lack metallic green
reflections) outside the virens/littorale group need to be sequenced to obtain a more robust
phylogeny and stabilize the classification. The name Virenshalictus Pesenko, 2007, is available
if a subgeneric status is found apropriate for the group in future studies.
Belgian Journal of Entomology 105: 124 (2020)
Fig. 15. Semi stabilized sand dunes, natural habitat of Lasioglossum dorchini in Alexander Stream National Park
(March 2017).
Fig. 16. Semi stabilized sand dunes, natural habitat of Lasioglossum dorchini in Alexander Stream National Park
(March 2017).
PAULY et al. A new species of bee from Israel
Fig. 17. Part of the Alexander Stream National Park planted with Eucalyptus trees (March 2017).
Fig. 18. Part of the Alexander Stream National Park planted with Eucalyptus trees (March 2018).
Belgian Journal of Entomology 105: 124 (2020)
The project HALAFREU was instrumental to the success of the present study, providing DNA analyses to support
the validity of the new species Lasioglossum dorchini and providing preliminary NJ-tree and phylogenetic tree for
the the virens/littorale group of species. This project was carried out by the Joint Experimental Molecular Unit
(JEMU), funded by the Belgian Science Policy. We thank Gideon Pisanty (SMNHTAU), Frank Koch and Viola
Richter (MNHUB), David Notton (NHML), and Andreas W. Ebmer (Linz) for the loan of types or museum
specimens of Lasioglossum leptocephalum and L. dorchini. Achick Dorchin (Tel Aviv University) found the types
of Lasioglossum sinaiticum at SMNHTAU and sent them to us on loan. He also captured the first specimen of the
new species dedicated to him. The following persons significantly helped us by recently collecting specimens of
the virens/littorale group of species: Pierre Rasmont (University of Mons, Belgium; specimens from Morocco),
Thomas J. Wood (Univeristy of Mons), Matilde Boschetti and Marino Quaranta (Italy), David Genoud, Alain
Livory and Eric Dufrêne (France).
The authors thank also the two referees, Tom Wood and Achik Dorchin, for numerous corrections and suggestions
to improve this paper, as well as Isabelle Coppée and late Isabelle Sauvage (SRBE) for reading the manuscript.
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Some types of unmanned aerial vehicles (UDs) are called drones. These equipment's of wide versatility have been applied very recently for different tasks, among which is agriculture. From there, this technology is responding to the decline of biopolynizers in different fields around the world. The objective of this work was to analyze the implications of the use of such devices in agricultural practices in the absence of entomophiles pollinators. Methodologically it was developed as a bibliographic review, consulting databases such as Scopus, Science Direct, Google academic and others; filtering by keywords, selecting the works according to the subject of interest and the criteria of the authors. Key information from the selected documents was extracted through the documentary analysis. As a result of this review, experiences related to the application of USNs in crop fertilization were found; the literature consulted also agrees to point out that drones are a viable strategy for such activity and is at the same time a solution to the diminution of entomophiles pollinators in certain regions of the Earth. However, the reference literature, very little reports possible adverse effects, which from an ecological point of view could arise from the use of USRs in agri-food production. It is concluded that further research is needed to understand the ecological consequences that may arise from the use of the equipment.
Full-text available
Morphological and allozyme analyses suggested the occurrence of a pseudocryptic species in the Lasioglossum villosulum (Kirby, 1802) species complex (Hymenoptera: Halictidae). We analysed the morphology of more than 1500 specimens and the DNA barcode fragment of the cytochrome c oxidase subunit I (COI) of 102 specimens of this species complex from several Palaearctic countries. Our phylogenetic tree reconstructions, based on maximum likelihood and Bayesian inference revealed one clade corresponding to all specimens morphologically identifi ed as Lasioglossum medinai (Vachal, 1895) and one divergent specimen morphologically identifi ed as Lasioglossum berberum (Benoist, 1941). The other specimens, morphologically identifi ed as L. villosulum, aggregated into at least three other lineages in our phylogenetic trees. The tree-based species delineations methods based on the Generalized Mixed Yule Coalescent (GMYC) model and the Bayesian Poisson Tree Process (bPTP) identifi ed fi ve to ten candidate species within the L. villosulum species complex, with L. medinai and L. berberum consistently recognized as separated from all other candidate species. Diagnostic morphological differences were found among L. medinai, L. berberum and the remaining specimens identified as L. villosulum. No diagnostic morphological differences were found to distinguish the different phylogenetic candidate species or lineages found within L. villosulum and L. medinai. Thus, both genetic and morphological approaches support the existence of L. medinai and L. berberum as distinct species from L. villosulum.
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Several forms or variants have long been recognized in the West Palearctic sweat bee Seladonia smaragdula (Vachal, 1895). Using DNA barcoding and morphological characters, primarily of the male genitalia, these variants are here recognized and described as five new species: S. gemmella Pauly sp. nov., S. submediterranea Pauly sp. nov., S. orientana Pauly & Devalez sp. nov., S. phryganica Pauly & Devalez sp. nov., and S. cretella Pauly & Devalez sp. nov. Also, we designate a lectotype for Halictus smaragdulus Vachal, consider Seladonia butea (Warncke, 1975) and S. morinella (Warncke, 1975) as nomina dubia, and discuss the identity of the Seladonia specimens from Australia currently determined as S. smaragdula.
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This study presents DNA barcode records for 4118 specimens representing 561 species of bees belonging to the six families of Apoidea (Andrenidae, Apidae, Colletidae, Halictidae, Megachilidae and Melittidae) found in Central Europe. These records provide fully compliant barcode sequences for 503 of the 571 bee species in the German fauna and partial sequences for 43 more. The barcode results are largely congruent with traditional taxonomy as only five closely allied pairs of species could not be discriminated by barcodes. As well, 90% of the species possessed sufficiently deep sequence divergence to be assigned to a different Barcode Index Number (BIN). In fact, 56 species (11%) were assigned to two or more BINs reflecting the high levels of intraspecific divergence among their component specimens. Fifty other species (9.7%) shared the same Barcode Index Number with one or more species, but most of these species belonged to a distinct barcode cluster within a particular BIN. The barcode data contributed to clarifying the status of nearly half the examined taxonomically problematic species of bees in the German fauna. Based on these results, the role of DNA barcoding as a tool for current and future taxonomic work is discussed. © 2015 The Authors. Molecular Ecology Resources Published by John Wiley Sons & Ltd.
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Phylogenies are increasingly used in all fields of medical and biological research. Moreover, because of the next generation sequencing revolution, datasets used for conducting phylogenetic analyses grow at an unprecedented pace. RAxML (Randomized Axelerated Maximum Likelihood) is a popular program for phylogenetic analyses of large datasets under maximum likelihood. Since the last RAxML paper in 2006, it has been continuously maintained and extended to accommodate the increasingly growing input datasets and to serve the needs of the user community. I present some of the most notable new features and extensions of RAxML, such as, a substantial extension of substitution models and supported data types, the introduction of SSE3, AVX, and AVX2 vector intrinsics, techniques for reducing the memory requirements of the code and a plethora of operations for conducting post-analyses on sets of trees. In addition, an up-to-date, 50 page user manual covering all new RAxML options is available. The code is available under GNU GPL at
There is an ongoing campaign to DNA barcode the world's >20,000 bee species. Recent revisions of Lasioglossum ( Dialictus ) (Hymenoptera: Halictidae) for Canada and the eastern United States were completed using integrative taxonomy. DNA barcode data from 110 species of L . ( Dialictus ) are examined for their value in identification and discovering additional taxonomic diversity. Specimen identification success was estimated using the best close match method. Error rates were 20% relative to current taxonomic understanding. Barcode Index Numbers (BINs) assigned using Refined Single Linkage Analysis (RESL) and 'barcode gaps' using the Automatic Barcode Gap Discovery (ABGD) method were also assessed. RESL is incongruent for 44.5% species, although some cryptic diversity may exist. Forty-three of 110 species were part of merged BINs with multiple species. The barcode gap is non-existent for the data set as a whole and ABGD showed levels of discordance similar to the RESL. The ' viridatum species-group' is particularly problematic, so that DNA barcodes alone would be misleading for species delimitation and specimen identification. Character-based methods using fixed nucleotide substitutions could improve specimen identification success in some cases. The use of DNA barcoding for species discovery for standard taxonomic practice in the absence of a well-defined 'barcode gap' is discussed.
A subgeneric classification for the 318 currently recognised species of the genus Evylaeus recorded from the Palaearctic Region (in the narrower understanding) is suggested. It includes 29 subgenera all of which are keyed. An annotated list of subgenera contains diagnoses and information about type species, general geographic distribution, and species included. The following 19 new subgenera are described: Aerathalictus subg. n. (type species: Melitta aerata Kirby), Biennilaeus subg. n. (type species: Halictus marginatus Brullé), Crassevylaeus subg. n. (type species: Halictus crassepunctatus Blüthgen), Fratevylaeus subg. n. (type species: Halictus fratellus Pérez), Glauchalictus subg. n. (type species: Halictus problematicus Blüthgen), Laevinodilaeus subg. n. (type species: Halictus laevinodis Morawitz), Limbevylaeus subg. n. (type species: Halictus limbellus Morawitz), Loethalictus subg. n. (type species: Halictus loetus Brullé), Malachevylaeus subg. n. (type species: Melitta malachura Kirby), Minutulaeus subg. n. (type species: Hylaeus minutulus Schenck), Monilevylaeus subg. n. (type species: Halictus immunitus Vachal), Nitidiusculaeus subg. n. (type species: Melitta nitidiuscula Kirby), Nodicornevylaeus subg. n. (type species: Halictus nodicornis Morawitz), Pallidevylaeus subg. n. (type species: Nomioides pallida Radoszkowski), Pauxevylaeus subg. n. (type species: Hylaeus pauxillus Schenck), Tricinctevylaeus subg. n. (type species: Halictus tricinctus Schenck), Truncevylaeus subg. n. (type species: Halictus truncaticollis Morawitz), Virenshalictus subg. n. (type species: Hylaeus virens Erichson), Viridihalictus subg. n. (type species: Halictus viridis Brullé). Hylaeus politus Schenck is fixed as the type species of the subgenus Pyghalictus Warncke 1975. Evylaeus albitarsoides (Blüthgen) is resurrected as a valid species. The following new synonymies are ascertained: Halictus fedtschenkoi Blüthgen 1938 = Lasioglossum andromeda Ebmer 1978, syn. n.; Lasioglossum fratellus betulae Ebmer 1978 = L. nupricola Sakagami 1988, syn. n. The lectotype of Halictus fedtschenkoi Blüthgen is designated. Advantages of a more finely divided and hierarchically deeper classification of large taxa are discussed. A synonymic catalogue of species-group names of the Palaearctic Evylaeus, comprising 679 names, appended.
Although commercial kits are available for automated DNA extraction, 'artisanal' protocols are not. In this study, we present a silica-based method that is sensitive, inexpensive and compliant with automation. The effectiveness of this protocol has now been tested on more than 5000 animal specimens with highly positive results. Ivanova). The present study seeks to overcome these constraints. Our study evaluated the effectiveness of several com- mercial glass fibre filtration (GF) plates, the core functional component in all silica-based DNA extraction kits. We worked only with 96-well plates that appeared structurally compatible with robotic systems. Specifically, we compared the performance of seven GF plates — two manufactured by BioLynx (B1, B2), three by PALL (P1-3) and two by Whatman (W1, W2). The B1 plate (no. F2008) was a 0.7- µ m glass fibre with 0.8 mL well; B2 (no. F2007) was a 1.0- µ m glass fibre with 0.8 mL well; P1 (no. 5051) was a 1.0- µ m glass fibre with 1 mL well; P2 (no. 5053) was a 3.0- µ m glass fibre media/0.2 µ m Bio-Inert membrane with 1 mL well; P3 (no. 5031) was a 1.0- µ m glass fibre media with 350 µ L well; W1 (no. 7505-0003) was a mini prep DNA binding plate with 0.8 mL well and W2 (no. 7700-7801) was a Unifilter plate with glass fibre media with 0.8 mL well. We began by identifying both buffer systems and protocols that enabled the use of these GF plates for manual DNA extraction. We then tested their performance under automation with a Biomek NX liquid-handling station (Beckman-Coulter) equipped with a filtration manifold. To provide a solid test of performance, we compared these results with those from a high performance commercial kit — the NucleoSpin96 (Machery-Nagel), hereafter termed the MN kit (Hajibabaei et al . 2005). Our performance comparisons employed DNA extracted from frozen tissues of six mammal species ( Glaucomys volans , Sorex fumeus , Clethrionomys gapperi , Blarina brevicauda,
The aims of this study are to review the current situation of the Israeli Mediterranean coastal sand dunes, to examine the causes for this situation, and to propose options for future conservation and management of the protected dune areas based on ecological, environmental, landscape and recreational demands and interests. The coastal dunes of Israel are characterized by diverse plant communities, with 173 plant species occurring on sand (8.2% of the total flora of Israel) including many endemic species (26% of all endemic species in Israel). Most of the species are annuals. The importance of the coastal strip as a centre of floral and faunal speciation is also manifested in the existing sand-bound animals. However, many species are rare. This is mainly due to the extensive industrial and urban development along the coastal plain and the direct and indirect destruction of the remaining open dune areas by tourism, recreation and sand mining. Only ca. 17% of the Israeli coastal dunes are still of good or reasonable ecological value, while < 5% of this area has been designated as protected area. Management policies differ from place to place and depend on local objectives. These objectives derive mainly from the knowledge and data that exist for each location, and its statutory status. Since 1995 several projects, which aim to develop integrated management tools for nature conservation and recreation uses for all coastal sand dunes in Israel have been conducted. These projects are summarized in the present paper.
In phylogenetic analyses of molecular sequence data, partitioning involves estimating independent models of molecular evolution for different sets of sites in a sequence alignment. Choosing an appropriate partitioning scheme is an important step in most analyses because it can affect the accuracy of phylogenetic reconstruction. Despite this, partitioning schemes are often chosen without explicit statistical justification. Here, we describe two new objective methods for the combined selection of best-fit partitioning schemes and nucleotide substitution models. These methods allow millions of partitioning schemes to be compared in realistic time frames and so permit the objective selection of partitioning schemes even for large multilocus DNA data sets. We demonstrate that these methods significantly outperform previous approaches, including both the ad hoc selection of partitioning schemes (e.g., partitioning by gene or codon position) and a recently proposed hierarchical clustering method. We have implemented these methods in an open-source program, PartitionFinder. This program allows users to select partitioning schemes and substitution models using a range of information-theoretic metrics (e.g., the Bayesian information criterion, akaike information criterion [AIC], and corrected AIC). We hope that PartitionFinder will encourage the objective selection of partitioning schemes and thus lead to improvements in phylogenetic analyses. PartitionFinder is written in Python and runs under Mac OSX 10.4 and above. The program, source code, and a detailed manual are freely available from