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JHR 69: I-21 (20 I 9) JOURNAL COR Pepeehrnieived open stents ocrost
doi: 10.3897/jhr.69.3256 | (4G) Hymenoptera

http://jhr.pensoft.net The Inerational Society of Hymenoptersts. RESEARCH

New genera of meliturguline bees from Saudi Arabia
and Persia, with notes on related genera and a key to
the Arabian fauna (Hymenoptera: Andrenidae)

Michael S. Engel'??, Abdulaziz S. Algarni*, Mohamed A. Shebl°,
Jennifer C. Thomas!

| Division of Entomology, Natural History Museum, 1501 Crestline Drive — Suite 140, University of Kansas,
Lawrence, Kansas 66045-4415, USA 2. Department of Ecology & Evolutionary Biology, University of Kansas,
Lawrence, Kansas 66045, USA 3 Division of Invertebrate Zoology, American Museum of Natural History,
Central Park West at 79” Street, New York, New York 10024-5192, USA 4 Department of Plant Protection,
College of Food and Agriculture Sciences, King Saud University, RO. Box 2460, Riyadh 11451, Saudi Arabia
5 Department of Plant Protection, Faculty of Agriculture, Suez Canal University, Ismailia, Egypt

Corresponding author: Michael S. Engel (msengel@ku.edu)

Academic editor: Michael Ohl | Received 19 December 2018 | Accepted 20 February 2019 | Published 30 April 2019
http://zoobank. org/5 B2D3ED0-DDC5-43A4-8564-AB 18373F1F9B

Citation: Engel MS, Algarni AS, Shebl MA, Thomas JC (2019) New genera of meliturguline bees from Saudi Arabia and
Persia, with notes on related genera and a key to the Arabian fauna (Hymenoptera: Andrenidae). Journal of Hymenoptera

Research 69: 1-21. https://doi.org/10.3897/jhr.69.32561

Abstract

A new genus of melitturgine bees (Panurginae: Melitturgini) is described and figured from central Saudi
Arabia. Belliturgula najdica Engel, gen. et sp. n., is similar in several respects to the genus Flavomeliturgula
Patiny in that both have a greatly elongate glossa that is longer than the face and is truncate apically. The
former differs in the greatly elongate and flattened first labial palpomere, the remaining palpomeres un-
modified, the greatly protuberant clypeus, and long labrum lacking dense setal patches, among other traits.
In addition, remarks are made on the composition of Flavomeliturgula, with Meliturgula deserta Warncke
(Iran, Pakistan), removed to Khuzimelissa Engel, gen. n., resulting in the new combination Khuzimelissa de-
serta (Warncke), comb. n. Khuzimelissa differs from Flavomeliturgula by the unmodified labial palpomeres
(lacking the elongate first three labial palpomeres of the latter genus), the acute glossa (broadly truncate in
the latter), glossa shorter than head length (greater than head length in the latter), the abundant, white,
squamiform setae of the mesoscutum (sparse, erect setae in the latter), cleft pretarsal claws (simple in the

latter), presence of metasomal setal bands (lacking in the latter), and apically emarginate pygidial plate

Copyright Michael S. Engel et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC
BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

2 Michael S. Engel et al. / Journal of Hymenoptera Research 69: 1-21 (2019)

(acutely rounded in the latter). The species of Flavomeliturgula are organized into three subgenera, includ-
ing Koreshomelissa Engel, subgen. n., and Freyamelissa Engel, subgen. n., and these may eventually warrant
generic status once phylogenetic work on the tribe has been completed. A key to the Arabian genera of

Panurginae is provided, along with an augmented key to the genera of subtribe Meliturgulina.

Keywords
Anthophila, Apoidea, Arabia, Melitturgini, Meliturgulina, Panurginae, taxonomy

Introduction

Bees of the panurgine tribe Melitturgini are uncommonly encountered in the Arabian
fauna. Hitherto, the tribe has been represented in Arabia by only four species in two
genera: Meliturgula scriptifrons (Walker) (=Poecilomelitta dzheddaensis Popov), M. ornata
(Popov), Borgatomelissa brevipennis (Walker) (= Nomia eburneifrons Walker, M. arabica
Popov), and B. niveopilosa Patiny (Walker 1871; Popov 1951; Patiny 2002). The biology
of these bees in Arabia remains unstudied, and even basic records of floral associations
are lacking. The latter two species have been considered related to Melitturga Latreille
and segregated into a nominate subtribe, while the species of Meliturgula Friese are clas-
sified in the subtribe Meliturgulina. Three genera have been recognized in Meliturgulina:
Gasparinahla Patiny, known only from a single, poorly preserved male from southwest-
ern Iran; Flavomeliturgula Patiny, encompassing several species occurring across southern
Iran and Pakistan; and Meliturgula, the largest genus of the group, with species distribut-
ed throughout Arabia and Africa but concentrated in southern Africa (Michener 2007).

Here we report a new species of meliturguline for the Arabian fauna, representing
a new genus. Individuals of the species were found collecting pollen from Carthamus
L. (Asteraceae: Carduoideae: Cynareae) in central Saudi Arabia (Figs 1, 2). The new
species is somewhat similar to some species hitherto classified in Flavomeliturgula. We
provide the description of the new taxon, along with notes on the systematics of Fla-
vomeliturgula (e.g., Table 1). The new species as well as several of those hitherto classi-

Table |. Current classification of species discussed herein.

Genus Khuzimelissa Engel, gen. n.

K. deserta (Warncke, 1983 [1985]) Iran (Khuzestan), Pakistan (Balochistan)
Genus Belliturgula Engel, gen. n.
B. najdica Engel, sp. n. Saudi Arabia (Riyadh)

Genus Flavomeliturgula Patiny, 1999
Subgenus Flavomeliturgula s.str.

E lacrymosa (Popov, 1967) Iran (Khuzestan, Hormozgan, Fars, Tehran)
Subgenus Freyamelissa Engel, subgen. n.

E schwarziana Patiny, 2004 Pakistan (Balochistan)

E berangeriae Patiny, 2002 Iran (Bushehr)
Subgenus Koreshomelissa Engel, subgen. n.

E tapana (Warncke, 1983 [1985]) Iran (Mazandaran)

E centaurea (Warncke, 1983 [1985]) Iran (Hormozgan), Pakistan (Balochistan)

New genera of meliturguline bees from Saudi Arabia and Persia, with notes on... 3

Figures |, 2. Female of Belliturgula najdica Engel, gen. et sp. n., at flowers of Carthamus L. (Asteraceae)

in central Saudi Arabia.

fied in Flavomeliturgula are as different from one another as are traditionally differenti-
ated as distinct genera among bees, particularly among different lineages of Panurginae
(e.g., Ruz 1986, 1991; Patiny 2001; Roig-Alsina and Compagnucci 2003; Gonzalez
and Engel 2011; Gonzalez et al. 2013, 2017; Ramos and Rozen 2014). Aside from
differences in setation, coloration, and the shape of labra, sulci, pygidial plates, and
other structural distinctions, the development of the mouthparts are strikingly differ-
ent among these groups, and this may reflect some degree of difference in host plant
visitation or specialization, much as is observed among other panurgine genera (e.g.,
Rozen and Ruz 1995; Rozen 2003). Accordingly, the classification is augmented in
order to reflect and highlight these differences, and encourage melittologists to see out
the biology of these rather different groups. In addition, we provide keys to the genera
of Arabian Panurginae and to the genera of subtribe Meliturgulina.

Material and methods

The morphological terminology for the systematic work is largely taken from Engel
(2001) and Michener (2007). Photographs were taken with a Canon 7D digital cam-
era attached to an Infinity K-2 long-distance microscope lens, illuminated with a Xe-
non flash, while measurements were taken with an ocular micrometer on an Olympus
SZX12 stereomicroscope. Specimens were examined from the collections of the Divi-
sion of Entomology (Snow Entomological Collections), University of Kansas Natural

4 Michael S. Engel et al. / Journal of Hymenoptera Research 69: 1-21 (2019)

History Museum, Lawrence, Kansas, USA (SEMC); the King Saud University Mu-
seum of Arthropods, Plant Protection Department, College of Food and Agriculture
Sciences, King Saud University, Riyadh, Kingdom of Saudi Arabia (KSMA); and the
Division of Invertebrate Zoology, American Museum of Natural History, New York,
New York, USA; while types for the new species described herein are deposited in the

former two institutions.

Systematics

Tribe Melitturgini Newman, 1834
Subtribe Meliturgulina Engel, 2001

Belliturgula Engel, gen. n.
http://zoobank.org/896747C8-0BD5-4B8B-8 DBA-3C33E1CC588A

Type species. Belliturgula najdica Engel, sp. n.

Diagnosis. The genus resembles Flavomeliturgula in the elongate and apically
truncate glossa but differs in the following attributes: body almost entirely pale yellow,
with scattered dark brown to black markings; outer subantennal sulcus arched; clypeus
greatly protuberant and extending well in front of compound eyes; labrum as long
as wide, with distinct basilateral ovoid windows of translucent integument; proximal
labial palpomere greatly elongate and flattened, twice as long as combined length of
remaining palpomeres, extending well past apex of paraglossa; second through fourth
labial palpomeres cylindrical, not flattened; apex of glossa not broadened; galea longer
than stipes; mesoscutum covered with abundant, erect, minutely spicate, white setae;
forewing Irs-m confluent with (rather than distad) 1m-cu; basal area of propodeum se-
tose; probasitarsus short and broad, about twice as long as wide; mesotibial spur short,
only about 0.5x length of mesobasitarsus; pygidial plate with margins concave apically
and therefore more abruptly tapering to acutely rounded apex.

Description. Small species (less than 8 mm in total length); integument largely
yellow, with some dark brown to black markings (Figs 1-5); head broader than long;
compound eyes with inner margins parallel (Fig. 5); outer subantennal sulcus greatly
arched outward (Fig. 5) (not straight); facial fovea shallow, narrower than median ocellar
diameter, elongate, length approximately 5.5x width (Fig. 5); clypeus greatly protuber-
ant, extending in front of compound eye by almost as much as compound eye width
(Fig. 8) (protuberance extending in front of compound eye by distance approximately
0.85—0.95x compound eye width); labrum as long as wide, with basilateral ovoid areas
(‘windows’) of translucent integument, without basilateral patches of dense, elongate,
branched setae, with shallow, mediolongitudinal depression; ventral surface of premen-
tum without abundant, long, posteriorly directed, simple setae (such setae present in
Khuzimelissa); first labial palpomere greatly elongate and flattened, much longer than
remainder of palpus (Fig. 12) (2x as long as combined lengths of remaining palpomer-

New genera of meliturguline bees from Saudi Arabia and Persia, with notes on... 5

Figures 3, 4. Female of Belliturgula najdica Engel, gen. et sp. n., from central Saudi Arabia. 3 Lateral
habitus 4 dorsal habitus.

es), extending greatly beyond apex of paraglossa, second palpomere not flattened, third
through fourth palpomeres short, subequal in length, cylindrical (Fig. 12); glossa greatly
elongate, distinctly longer than face (Fig. 8); apex truncate but not broadened (Fig. 9);
galea longer than stipes (galea slightly longer than stipes in Flavomeliturgula; galea shorter
than stipes in Khuzimelissa); mesoscutum with numerous scattered, erect, minutely spi-
cate (pipecleaner-like in appearance), white setae, not obscuring integument (0.8—1.3x

6 Michael S. Engel et al. / Journal of Hymenoptera Research 69: 1-21 (2019)

Figures 5-7. Female faces of representative Meliturgulina. 5 Belliturgula najdica Engel, gen. et sp. n.,

from Saudi Arabia (note arched outer subantennal sulci) 6 Khuzimelissa deserta (Warncke), comb. n.,
from Pakistan 7 Flavomeliturgula (Flavomeliturgula) lacrymosa (Popov) from Iran (note straight outer
subantennal sulci).

median ocellar diameter) (Fig. 4); basal area of propodeum setose, setae not obscuring
integument; forewing with pterostigma longer than wide, approximately 3.5x as long as
wide, width subequal to length of prestigma, about 3.5x as long as prestigma, margin bor-
dering first submarginal cell slightly diverging from anterior wing margin, border inside
marginal cell converging toward anterior wing margin and with convex arch; marginal
cell widely and slightly obliquely truncate at apex, longer than distance from its apex to
wing tip; three submarginal cells present (i.e., 1rs-m present); length of first submarginal
cell slightly greater than combined lengths of second and third submarginal cells; r-rs

New genera of meliturguline bees from Saudi Arabia and Persia, with notes on... 7

and 3Rs (anterior border of second submarginal cell) subequal; 2Rs and Irs-m slightly
converging anteriorly, thus second submarginal cell trapezoidal; 3Rs slightly longer than
0.5x 4Rs (anterior border of third submarginal cell); 1rs-m confluent with 1m-cu (as
in Gasparinhala); \cu-a at least 2x as long as 2M+Cu (usually 2M+Cu even shorter
in other genera); hind wing with six distal hamuli; probasitarsus broad, about twice as
long as wide (probasitarsus more elongate slender in Flavomeliturgula, nearly four times
as long as wide); protibial calcar with velum rectangular, velum with outer and apical
borders comparatively straight, malus (apical portion of rachis: sensu Engel et al., 2017)
ciliate and slightly shorter than velum; mesofemur ventro-proximally forming nearly
orthogonal angle (as in Flavomeliturgula, Meliturgula, Khuzimelissa, and related genera);
mesotibial spur serrate, about 0.5x length of mesobasitarsus; metabasitibial plate broadly
rounded, with carinate margins and surface covered in appressed, minute, plumose setae;
metatibial and metabasitarsal scopa composed of abundant, elongate (1.8—3.2x median
ocellar diameter), simple setae; pretarsal claws simple; metasomal terga without well-de-
veloped setal bands (Fig. 4) (present in Khuzimelissa); pygidial plate with apex narrowly
rounded, margins tapering more prominently in apical half (Fig. 24).

Etymology. The generic name is an artificial combination of Bell, honoring explor-
er, archeologist, and nation-builder Gertrude Margaret Lowthian Bell (1868-1926),
and a subset of Meliturgula Friese, type genus of the subtribe Meliturgulina. The name
is therefore treated as a euphonious combination of letters and the gender of the name
is considered to be feminine.

Included species. The genus includes at present only the type species, found in
central Saudi Arabia.

Belliturgula najdica Engel, sp. n.
http://zoobank.org/145B107D-91C2-424D-A736-5521CD3FF7DD
Figs 1-5, 8, 9, 12, 24

Diagnosis. As for the genus (vide supra).

Description. 9: Total body length 7.08-7.33 mm; forewing length 4.13—
4.27 mm. Head broader than long, width 2.08-2.13 mm, length 1.67—1.71 mm;
compound eyes with inner orbits margins roughly parallel, upper interorbital dis-
tance 1.33-1.38 mm, lower interorbital distance 1.33—1.36 mm; distance between
compound eyes slightly greater than compound eye length; compound eye length
1.25 mm, width 0.58—0.63 mm; clypeus greatly protuberant, extending in front of
compound eye by nearly compound eye width; intertegular distance 1.42—1.46 mm.

Clypeus and supraclypeal area largely smooth with coarse, shallow (almost faint in
most areas) punctures, such punctures separated by a puncture width or more, slightly
denser along borders; labrum similar to clypeus except impunctate on transparent ovals;
remainder of face similar except punctures smaller, more defined (albeit still shallow),
and separated by a puncture width or frequently less; dark area between ocelli minutely
and faintly imbricate and impunctate; facial fovea imbricate and impunctate; gena as on

8 Michael S. Engel et al. / Journal of Hymenoptera Research 69: 1-21 (2019)

Figures 8-11. Female heads and mouthparts in profile and details of glossal apexes. 8 Head profile of
Belliturgula najdica Engel, gen. et sp. n., from Saudi Arabia 9 glossal apex of B. najdica 10 head profile
of Flavomeliturgula (Flavomeliturgula) lacrymosa (Popov) from Iran II glossal apex of F (F) lacrymosa.

upper face except punctures becoming more sparsely separated ventrally; mesoscutum
smooth to faintly and minutely imbricate with shallow punctures separated by a punc-
ture width or less; mesoscutellum as on mesoscutum except punctures slightly denser
and integument slightly more noticeably imbricate; metanotum as on mesoscutellum
except shallow punctures nearly contiguous; pleura as on mesoscutum except punctures
shallower and becoming sparse ventrally; basal area of propodeum as on metanotum;

New genera of meliturguline bees from Saudi Arabia and Persia, with notes on... )

lateral and posterior surfaces of propodeum with small punctures separated by a punc-
ture width or frequently less, integument between punctures smooth to faintly imbri-
cate; metasomal terga minutely and finely imbricate with scattered shallow, faint, small
punctures separated by a puncture width or less, such punctures most easily discernible
on first tergum and becoming fainter on succeeding terga; pygidial plate minutely and
finely imbricate; sterna as on terga except shallow, coarse punctures more prominent.

Integument generally pale yellow except marked with dark brown to black on
mandibular apex, on facial fovea, around ocelli (Figs 4, 5); mesoscutum with three lon-
gitudinal dark stripes, medial stripe thin, terminating before posterior of mesoscutum,
lateral stripes thicker, situated between lateral yellow borders and medial stripe, ex-
tending to axilla (Fig. 4); dark brown on lateral depression of axilla and along anterior
border of mesoscutellum (slightly extended mediolongitudinally) (Fig. 4), and light
brown anteriorly on basal area of propodeum (Fig. 4). Wing membranes clear, hyaline;
veins pale yellow to off white (Fig. 3). Metasoma with light to dark brown marks as fol-
lows: paramedian spots on anterior-facing surface of first metasomal tergum (Fig. 4),
elongate oval spots at lateral extremities of second tergum (Fig. 3), small spots on lat-
eral extremities of third tergum (absent in some individuals); terga I-V with irregular
brownish-purplish coloration (reflective of inner organs through pale integument) to
varying degrees (in some individuals inner gut prominent through integument, others
less so); pygidial plate brown, becoming reddish apically (Fig. 24); sterna pale brown,
becoming darker on two apical sterna.

Pubescence generally white and largely sparse (Figs 3-5); facial setae generally short
to moderate (0.2—1.0x median ocellar diameter) in length and minutely spicate (i.e.,
minutely branched on all sides along entire length of rachis, with a pipecleaner-like ap-
pearance), such setae scattered on face and generally suberect; scape with similar moder-
ate-length, erect, minutely spicate setae; minutely spicate setae longer (0.9—1.5x median
ocellar diameter) and erect around ocelli, vertex, posterior of gena, and postgena; man-
dible with more sparse, elongate (1.8—3.0x median ocellar diameter), simple setae along
lower margin; mesoscutum with numerous erect short to moderate-length (0.4—1.2x
median ocellar diameter) minutely spicate setae, such setae not obscuring integument;
mesoscutellum as on mesoscutum although setae slightly sparser; metanotum with mi-
nutely spicate setae sparse on disc, with various greatly elongate (up to 2.0x median
ocellar diameter) minutely spicate setae at lateral extremities; pleura with numerous
moderate-length to elongate (1.0—2.2x median ocellar diameter) minutely spicate setae,
such setae not obscuring integument; basal area of propodeum with numerous fine mi-
nutely spicate setae, not obscuring integument; propodeum with lateral and posterior
surfaces with pubescence similar to that of pleura; legs with minutely spicate setae ex-
cept simple on tarsi and scopal setae of metatibia and metabasitarsus elongate (1.8—3.2x
median ocellar diameter) and simple; metasoma with generally sparse pubescence, setae
more numerous on apical terga where such setae minutely spicate and suberect to erect;
setae particularly dense around pygidial plate and obscuring integument.

CS: Latet.

10 Michael S. Engel et al. / Journal of Hymenoptera Research 69: 1-21 (2019)

er a aS 5 Re
\

wie | Ee

0.5mm

Figures 12-15. Labial palpi of representative species (to same scale) and labiomaxillary complex of
Khuzimelissa deserta (Warncke), comb. n. (not to same scale), all in ventral view. 12 Labial palpus of Be/-
liturgula najdica Engel, gen. et sp. n. 13 labial palpus of K. deserta 14 labial palpus of Flavomeliturgula
(Flavomeliturgula) lacrymosa (Popov) |5 photograph of labiomaxillary complex of K. deserta.

Holotype. 2, KSA [Kingdom of Saudi Arabia], Riyadh [Region], Derab Agricul-
tural Research Station [24°25'23"N, 46°39'01"E, 580 m elevation], 29.4.2015 [29
April 2015], M. Shebl, Carthamus |Asteraceae: Carthamus L.] (SEMC).

Paratypes. 2° 9, same data as holotype (SEMC, KSMA).

Etymology. The specific epithet is taken from the Najd, the vast geographic central
plateau portion of Saudi Arabia which encompasses the type locality which is slightly
south of the city of Riyadh.

Comments. The type series was collected from an unidentified species of distaff
thistle (Asteraceae: Carduoideae: Carthamus L.) growing around the Dirab Agricultural
Research Station in central Saudi Arabia (just south from Riyadh). The most common
species of Carthamus in Saudi Arabia are Carthamus oxyacantha M.Bieb., which grows
wild in many disturbed areas, and C. tinctorius Linnaeus (safflower), the latter widely
cultivated, particularly so in southern Arabia (Mandaville, 1990). It is likely that the
type series of B. najdica was captured at C. oxyacantha. Species of Carthamus are spiny,
dentate herbs that can be largely self-pollinated, although among insects bees are the
dominant pollinators (e.g., Butler et al., 1966).

New genera of meliturguline bees from Saudi Arabia and Persia, with notes on... 11

Khuzimelissa Engel, gen. n.

http://zoobank.org/FF287D5A-5 1 DA-4BE4-A353-B1B896BB858D

Type species. Meliturgula deserta Warncke, 1983 [1985].

Diagnosis. The genus superficially resembles Flavomeliturgula and Belliturgula,
particularly the latter in the extensive yellow coloration. However, the genus differs
from both in the short, more typical labial palpomeres in which the proximal labial
palpomere is elongate, flattened, longer than the combined lengths of the remaining
palpomeres (approximately 1.25x length of remaining palpomeres combined), with
the second palpomere flattened and slightly shorter than the length of the third and
fourth palpomeres combined; and the third and fourth palpomeres subequal in length
and cylindrical (Fig. 13). In addition, the glossa is acute (Figs 6, 15) (rather than
broadly truncate apically), the glossa is shorter than the head length (Figs 6, 19-21)
(rather than greater than the head length in the former two genera), the pretarsal
claws are cleft (simple in the former two genera), the metasomal terga have distinctive

broad setal bands (Figs 16-18) (lacking in the former two genera), and the pygidial

Figures 16-18. Female of Khuzimelissa deserta (Warncke), comb. n., from Pakistan. 16 Dorsal habitus
17 lateral habitus 18 detail of mesosoma and anterior metasomal dorsum.

2 Michael S. Engel et al. / Journal of Hymenoptera Research 69: 1-21 (2019)

Figures 19-21. Female of Khuzimelissa deserta (Warncke), comb. n., from Pakistan. 19 Head and
mouthparts in profile 20 dorsal-frontal view of mouthparts 21 extended labiomaxillary complex in profile.

plate is apically emarginate (Fig. 25) (acutely rounded in the former two genera).
Other diagnostic traits include: small species, slightly more than 8 mm in length; in-
tegument largely pale yellow with few dark brown to black markings (Figs 6, 16-18);
outer subantennal sulcus arched (Fig. 6); clypeus gently and weakly convex, scarcely
protuberant (Figs 6, 17, 19), extending in front of compound eye by about 0.5x com-

New genera of meliturguline bees from Saudi Arabia and Persia, with notes on... i

Figures 22, 23. Female of Flavomeliturgula (Flavomeliturgula) lacrymosa (Popov) from Iran. 22 Dorsal
habitus 23 lateral habitus.

pound eye width; labrum wider than long, width 1.7x length, without lateral oval
windows of transparent integument (Fig. 20), surface gently and weakly convex with
weak subtriangular proximal medial depression; prementum ventrally with abundant,
elongate (1.0—1.9x median ocellar diameter), erect to suberect, simple setae (Figs 15,
19, 21); galea shorter than stipes; mesoscutum with abundant, appressed, squamose
setae (Fig. 18), intermixed with sparse, erect, minutely spicate setae (0.8—1.0x medi-
an ocellar diameter); probasitarsus elongate, slender, length approximately 3x width;
mesotibial spur about as long as mesobasitarsus; forewing with r-rs distinctly shorter

14 Michael S. Engel et al. / Journal of Hymenoptera Research 69: 1-21 (2019)

Figures 24-26. Female pygidial plates. 24 Belliturgula najdica Engel, gen. et sp. n. 25 Khuzimelissa
deserta (Warncke), comb. n. 26 Flavomeliturgula (Flavomeliturgula) lacrymosa (Popov).

than 3Rs, Irs-m distad 1m-cu, and Icu-a about 4x as long as 2M+Cu; hind wing with
10 or more distal hamuli arranged in a dense series.

Etymology. The new genus-group name is a combination of Khuzi, the Middle
Persian name for the ancient Susian people, and from which the regional name Khuz-
estan derives (and encompasses the type locality of the type species), and the Greek,
mélissa (usitoou), meaning, “bee”. The gender of the name is feminine.

Included species. ‘The genus presently includes only the type species, Khuzimelissa
deserta (Warncke), comb. n., which has been recorded from Iran (Khuzestan) and Pa-

kistan (Balochistan).
Genus Flavomeliturgula Patiny
Diagnosis. The genus can be generally differentiated from other members of the

subtribe by the following combination of features: small to medium-sized bees, 5.0—
10.0 mm in length; integument dark brown to brown with extensive yellow mark-

New genera of meliturguline bees from Saudi Arabia and Persia, with notes on... 15

ings (Figs 22, 23); outer subantennal sulcus straight (Fig. 7); clypeus weakly protuber-
ant, extending in front of compound eye by about 0.35—0.5x compound eye width
(Fig. 10); labrum wider than long, width 1.5x length, without lateral oval windows of
transparent integument, surface with weakly raised, medial plateau, basilaterally with
prominent setose patches composed of long (1.8—2.4x median ocellar diameter), sub-
decumbent setae; prementum ventrally without elongate setae; proximal labial pal-
pomere elongate (but not extending past apex of paraglossa: in Belliturgula palpomere
extends well past apex of paraglossa), flattened, about as long as to slightly longer than
combined lengths of remaining palpomeres; second and third palpomeres flattened
and elongate, together slightly shorter than to about as long as first palpomere, fourth
palpomere short, shorter than preceding palpomere, cylindrical combined (Fig. 14);
glossa greatly elongate, longer than head length, apex broadly truncate and broadened
(Figs 7, 10, 11); mesoscutum without appressed, squamose setae or abundant spicate
setae (Fig. 22); forewing with 1rs-m distad 1m -cu (confluent in Belliturgula); proba-
sitarsus elongate, slender, length approximately 4x width; mesotibial spur elongate,
nearly 0.75x as long as mesobasitarsus; pretarsal claws simple; metasomal terga with-
out distinctive broad setal bands; pygidial plate apically acutely rounded, with lateral
margins tapering evenly toward apex, surface with mediolongitudinal ridge (Fig. 26).

Key to Subgenera of Flavomeliturgula

1 Propodeum basal area without setae; hind wing with 10 or more distal ha-
muli arranged in a dense series ..............1e Flavomeliturgula Patiny, s.str.
= Propodeum basal area with distinct setae; hind wing with 6—7 distal hamuli.
aR ot ee gatae 2) iG Meats Re Naas GI NRF ce eree Sn Se Acar Re oan ae Re ane pA
2(1) Mesoscutum brown to black with yellow markings; mesoscutum with min-

ute, scattered setae, length of setae distinctly less than 0.5x median ocellar
GTO ET has hoes tas er ean tes Beene au bel Koreshomelissa Engel, subgen. n.
= Mesoscutum entirely black, without yellow markings; mesoscutum with scat-
tered, white setae, particularly anteriorly, length of setae 1—-1.5x median ocel-
lars diainieters.c.0F cseetessite thd tn Ueeteteete toe. Freyamelissa Engel, subgen. n.

Subgenus Flavomeliturgula Patiny

Meliturgula (Flavomeliturgula) Warncke, 1983 [1985]: 229, nomen invalidum (no type
species designated: ICZN 1999: Art. 13.3). Nota bene: Michener (2007) errone-
ously indicated the name as used by Warncke was a nomen nudum and included
no species. In fact, Warncke did include four species and even provided scant diag-
nostic traits for the group (meaning it was not a nomen nudum), but only failed to
explicitly designate a type species.

16 Michael S. Engel et al. / Journal of Hymenoptera Research 69: 1-21 (2019)

Flavomeliturgula Patiny, 1999: 251. Type species: Poecilomelitta lacrymosa Popov, 1967,
by original designation.

Diagnosis. The subgenus can be distinguished by the combination of: mesoscutum
dark brown to black with yellow markings, surface with scattered, appressed, fine se-
tae, such setae mostly anteriorly and laterally, and 0.5—0.75x median ocellar diameter;
basal area of propodeum without setae; and hind wing with 10 or more distal hamuli
arranged in a dense series along anterior wing margin.

Included species. The subgenus as here circumscribed is restricted to the type species,
Flavomeliturgula lacrymosa (Popov), from Iran (Fars, Hormozgan, Khuzestan, Tehran).

Koreshomelissa Engel, subgen. n.

http://zoobank.org/9A888F48-0556-4243-A804-07000235BB63

Type species. Meliturgula tapana Warncke, 1983 [1985].

Diagnosis. The subgenus can be distinguished by the combination of: mesoscu-
tum dark brown to black with yellow markings, surface with scattered, minute, fine
setae, and less than 0.5x median ocellar diameter; basal area of propodeum with setae;
and hind wing with 6-7 distal hamuli along anterior wing margin.

Etymology. The new subgeneric name is a combination of KéréS or Kurus, the
Hebrew and Persian names for Cyrus the Great (Cyrus II, ca. 600-530 B.C.), who first
unified the ancient kingdoms of the Near East and founded the Achaemenid Empire
of Persia, combined with the Greek, mélissa (uéioon), meaning, “bee”. The gender of
the name is feminine.

Included species. The subgenus includes two species: F (Koreshomelissa) tapana and
E (K.) centaurea (Warncke), both from northern (along the Caspian Sea) and southern
costal Iran (Mazandaran and Hormozgan) and southern Pakistan (Balochistan).

Freyamelissa Engel, subgen. n.

http://zoobank.org/6F683 DEC-CFBB-4FFA-B998-9D4333967A4B

Type species. Flavomeliturgula berangeriae Patiny, 2002.

Diagnosis. The subgenus can be distinguished by the combination of: mesoscu-
tum entirely black without yellow maculation, surface with scattered setae of 1-1.5x
median ocellar diameter; basal area of propodeum with setae; and hind wing with 6—7
distal hamuli along anterior wing margin.

Etymology. The new subgeneric name is a combination of Freya, for the famed
explorer of Arabia, Persia, and Afghanistan, Freya Madeline Stark (1893-1993), and
the Greek, mélissa (uéioox), meaning, “bee”. The gender of the name is feminine.

Included species. The subgenus includes two species: F (Freyamelissa) berangeriae
and F (F) schwarziana Patiny, from southwestern Iran (Bushehr) and southern Paki-
stan (Balochistan).

New genera of meliturguline bees from Saudi Arabia and Persia, with notes on... 17

Key to Genera of Saudi Arabian Panurginae

1

(1)

3(1)

4(3)

Forewing with two submarginal cells 0... eee esses ceseeseeeeeeseeseseeeeseeeeees 2
Forewing withsthree-submaarentra lee IS. 5183.0) Shancrubect to ahab ten viit hos nhayten sidoban olan 3
S6 of male with posterior margin thin and more or less straight or usually
broadly bilobed but usually without an apical zone of very short setae; epis-
ternal groove present but usually short, not reaching level of scrobe, or rarely
absent; 1m-cu distal to 1rs-m, joining second submarginal cell...

sSrsuinev Sainsten ake Sotny Meant go sine ti Pa reat One'seneyoreecavenieesiny Siren oe ester Paes spy tgs Panurgus Panzer
S6 of male with posterior margin slightly to strongly produced medially, this
projection with truncate to broadly concave apex margined by zone of short
setae; episternal groove entirely absent (body entirely black except for yellow
clypeus of some males; 1m-cu meeting or basal to 1rs-m or nearly so ...........

PhSNa assets at Rhee A Misa aes tae scsils ge th ee Panurginus Nylander
First labial palpomere greatly elongate and flattened, much longer than re-
mainder of palpus, third through fourth palpomeres short, subequal in
length, cylindrical; glossa longer than face, with apex truncate; body wholly
yellow with few black markings on head and mesosomal dorsum................4.

secs Sie bn on essen Ps eon Bion ie Cee ate a Belliturgula Engel, gen. n.
First labial palpomere elongate, as long as or longer than second to fourth
palpomeres together; glossa pointed; body variable mostly dark, never wholly
LLG clos os "Seey Pastcalar go aetna Pro's eat oo se psmanc'e enter spice vay Bree ov onsen ese oe sees Mic eden aeenerecenee 4
Mesoscutum of female densely covered with short, thickened, appressed setae
among which are scattered erect setae; male gonostylus very slender, as long as
gonocoxite; volsella spiculate, extending beyond gonocoxites; seventh meta-
somal tergum of male with bifid apex; sixth and seventh metasomal sterna of
male with deep medioapical emarginations............... Borgatomelissa Patiny
Mesoscutum of female with scattered suberect setae or in some species with
appressed setae, less dense than as above; male gonostylus robust, half as long
as gonocoxite or less; volsella not spiculate, extending little if at all beyond
gonocoxites; seventh metasomal tergum of male with apex simple; sixth and
seventh metasomal sterna of male with apical margins transverse or convex..

be PR ee a P irc ite i cr eer RO ny OR OR A AO Meliturgula Friese
Key to Genera of Meliturgulina
1 Glossarshortemthansiace’... scree occa cece Se ae 2
- Glossatloncermthansace sta metus. Merete oa eta sheer unter taee tel ak ratens teal oak 3
21) Body largely yellow; pygidial plate of female emarginate apically, surface with

weak mediolongitudinal ridge, faintly imbricate 0.0... lee eee eeeeeeeeeeeeees
sansa hadi txt AN ted aca thins Land Ga Ms ie AA a Khuzimelissa Engel, gen. n.
Body largely black with areas of yellow maculation; pygidial plate of female
rounded apically, surface flat, imbricate ....... eee Meliturgula Friese

18 Michael S. Engel et al. / Journal of Hymenoptera Research 69: 1-21 (2019)

Sl) Glossal apex broadly truncate; maxillary palpus with six palpomeres........... 4
- Glossal apex acute; maxillary palpus apparently vestigial (“nearly absent” ac-

COrding 16-Pariiiy, DOU) wccciercocerscesctetsonntisaieeticetes Gasparinahla Patiny
4(3) Proximal labial palpomere greatly elongate, more than 2x as long as com-

bined lengths of remaining palpomeres, extending greatly beyond apex of
paraglossa; remaining palpomeres not flattened; outer subantennal sulcus of
female greatly arched; female clypeus greatly protuberant, in profile extend-
ing in front of compound eye by about compound eye width; female labrum
as long as wide, without medioapical elevated plateau and without basilateral
patches of dense, long, branched setae ............... Belliturgula Engel, gen. n.
- Proximal three labial palpomeres long, flattened; first labial palpomere not ex-
tending past apex of paraglossa, shorter than combined lengths of remaining
palpomeres; outer subantennal sulcus straight; clypeus weakly protuberant,
in profile extending in front of compound eye by much less than compound
eye width; female labrum wider than long, with medioapical elevated circular
or ovoid plateau, with basilateral patches of dense, long, branched setae.......

igh sok petra ss Ae A Mar PRE eh neg scteseal Baek re ae Flavomeliturgula Patiny

Discussion

The diversity of bee genera occurring across the Arabian Peninsula has been summarized
most recently by Dathe (2010) and Engel et al. (2013, 2017). In recent years several
genera have been added to the Arabian fauna, including Melitta Kirby (Melittidae),
Compsomelissa Alfken, and Tarsalia Morawitz (both Apidae) (Algarni et al., 2014; Shebl
et al., 2016; Engel et al., 2014, 2017). Prior to the present work, 45 genera of bees have
been documented from Saudi Arabia (Engel et al., 2017), and only four panurgines
(Borgatomelissa, Meliturgula, Panurginus Nylander, and Panurgus Panzer). None of these
previously documented genera are endemic to Arabia (e.g., Patiny and Michez 2007;
Michener 2007), and most of these genera are widely distributed across the Palaearctic
or even throughout the Old World. Here we record a unique member of the panurgine
tribe Melitturgini, representing a genus allied to a group of species largely occurring
across Persia and encompassing a variety of morphologies, as documented above. The
few known occurrences for the groups discussed herein (Table 1) are loosely analogous
to those of species in the genus 7arsalia (Apinae: Tarsaliini) (Engel et al., 2017). The sub-
genera and species of Tarsalia are distributed across Persia and the Middle East, but also
extend into western India and into Central Asia to the East and into Egypt and Sudan
in the West. Unfortunately, the biology of this group of bees, comprising the genera Bel-
liturgula, Flavomeliturgula, and Khuzimelissa, remains largely unknown aside from a few
floral records. Interestingly, like the new Arabian species described herein, other species
where records are available from Iran have also been found at flowers of the thistle sub-
family Carduoideae (Asteraceae), and specifically visiting the genera Centaurea L. and
Carthamus L. (both of tribe Cynareae) (Warncke, 1983). It is possible that all of these

New genera of meliturguline bees from Saudi Arabia and Persia, with notes on... 19

genera are oligolectic on genera of Carduoideae or more generally across Asteraceae.
It is greatly hoped that by drawing attention to these unique bees, and the interesting
mouthpart specializations found in each (e.g., glossal structure, forms of the labial palpi,
premental setation), that efforts will be made to seek out further material in the field,
obtain observations on their floral visitation behaviors, locate nests and immatures, and
discover the unknown males or females since many of these species are known from a
single sex. It is likely such exploration will also uncovered as of yet unknown species,
as well as document additional localities from which meaningful distributions can be
mapped, ecological niches determined, and broader biogeographic patterns revealed.
Naturally, a comprehensive phylogeny of meliturgulines would be of considerable value,
but at present the rarity of material (i.e., only one species of the group is known from
both sexes and from any substantial number of locality records and specimens) hinders
any meaningful study as characters would ideally be drawn from both males and fe-
males. Similarly, for DNA sequence data additional collections from the field are neces-
sary. Ultimately, more fieldwork is necessary across the regions in which these bees occur.

Acknowledgements

The authors extend their appreciation to the International Scientific Partnership Pro-
gram (ISPP) at King Saud University for funding this research through ISPP #0083.
The manuscript received important and helpful reviews from two anonymous evalua-
tors, to whom we are most grateful.

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