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Belgian Journal of Entomology 105: 124 (2020) ISSN: 2295-0214
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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
& Yael MANDELIK2
1 Royal Belgian Institute of natural Sciences, O.D. Taxonomy & Phylogeny, Rue Vautier 29, 1000 Brussels,
Belgium; e-mails: alain.pauly54@gmail.com, apauly@naturalsciences.be (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
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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
24.
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)
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Lasioglossum dorchini (Hymenoptera: Apoidea: Halictidae)
a new species of bee from Israel
Alain PAULY1, Karmit LEVY2, Grégoire NOËL3, Gontran SONET1, Jean-Luc BOEVÉ1
& Yael MANDELIK2
1 Royal Belgian Institute of natural Sciences, O.D. Taxonomy & Phylogeny, Rue Vautier 29, 1000 Brussels,
Belgium; e-mails: alain.pauly54@gmail.com, apauly@naturalsciences.be (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.
Abstract
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
Résumé
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.
Introduction
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
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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
REPOSITORIES AND EXAMINED SPECIMENS
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).
PROTOCOL FOR DNA ANALYSIS
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).
PHYLOGENETIC ANALYSIS
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)
5
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.
Results
Lasioglossum dorchini Pauly sp. nov.
urn:lsid:zoobank.org:act:9A1ED256-0C1F-497E-A7CF-C551C1049610
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).
DESCRIPTION. FEMALE. Length 6 mm.
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
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Table 1. Female specimen information from the virens / littorale group with their GenBank accession or BOLD
process ID.
Code
Species
Locality
Date
Leg.
AP221
Lasioglossum aureolum
France
2014
D. Genoud
AP303
Lasioglossum
pseudolittorale
Israel, Nahal Alexander
2016
K. Levy
AP304
Lasioglossum dorchini
Israel, Nahal Alexander
2016
K. Levy
AP404
Lasioglossum aureolum
Portugal, Tavira, Cabanas
2016
A. Livory & R.
Coulomb
AP409
Lasioglossum aff littorale
Portugal, Barra
2016
A. Livory & R.
Coulomb
AP410
Lasioglossum littorale
littorale
Italy, Toscana, Parco regional
MSRM
(Migliarino san Rossore
Massaciucolli)
2016
M. Boschetti
AP454
Lasioglossum
pseudoleptocephalum
Portugal, Algarve, Praia do Barril,
Tavira
2016
T. Wood
AP463
Lasioglossum aff littorale
Italy, Sardarnia, Cagliari,
Castiadas, San Giusta
2015
E. Dufrêne
AP499
Lasioglossum
leptocephalum
Israel, Park Britannia
2016
T. Chaprazaro
AP500
Lasioglossum dorchini
Israel, Nahal Alexander
2016
K. Levy
AP501
Lasioglossum dorchini
Israel, Nahal Alexander
2016
K. Levy
AP502
Lasioglossum dorchini
Israel, Nahal Alexander
2017
K. Levy
AP503
Lasioglossum dorchini
Israel, Nahal Alexander
2018
K. Levy
AP699
Lasioglossum
leptocephalum
Israel, Holot Shunera
2015
G. Pisanty
AP700
Lasioglossum
leptocephalum
Israel, Holot Shunera
2015
G. Pisanty
AP701
Lasioglossum musculoides
Morocco, Haddada
2018
I. El Abdouni
AP702
Lasioglossum aff littorale
Morocco, Bouknadel
2018
I. El Abdouni & P.
Lhomme
AP703
Lasioglossum aff littorale
Morocco, Haddada
2018
I. El Abdouni & P.
Lhomme
AP704
Lasioglossum aff littorale
Morocco, Bouknadel
2018
I. El Abdouni & L.
Harroud
AP705
Lasioglossum aff littorale
Morocco, Haddada
2018
I. El Abdouni & A.
Sentil
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)
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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
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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)
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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
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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)
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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
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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)
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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
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TYPE MATERIAL (456 specimens; SMNHTAU, RBINS). Holotype : ISRAEL, Nahal
Alexander, 32.401 34.882, 1.V.2016, leg. K. Levy (AP 304) (GenBank MT221620)
(SMNHTAU).
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)
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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
16
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)
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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)
(SMNHTAU).
PAULY et al. A new species of bee from Israel
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Fig. 12. Distribution map of Lasioglossum leptocephalum.
Table 2. Morphological comparison between Lasioglossum leptocephalum and L. dorchini
Characters and measurements
Lasioglossum leptocephalum
Lasioglossum dorchini
Female
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)
Male
(type of Halictus krugeri
Blüthgen 1930)
(males from Nahal
Alexander)
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.
11c)
wrinkles reaching posterior
margin of propodeal area
(Fig. 8c).
Belgian Journal of Entomology 105: 124 (2020)
19
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
20
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)
21
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
22
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)
23
Acknowledgements
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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