European Journal of Taxonomy 174: 1-18

http://dx.doi.org/10.5852/ejt.2016.174

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ISSN 2118-9773

www. europeanj ournaloftaxonomy. eu

2016 • Litman J.R. et al.

Research article

urn:lsid:zoobank.org:pub:DOB 13968-516F-4QC4-9C0B-4C4DCF362D13

A new species of Fidelia Friese, 1899 (Hymenoptera, Megachilidae),

with a key to the species of the genus

Jessica R. LITMAN '•*, Connal D. EARDLEY 2 & Michael KUHLMANN 3

1 Natural History Museum of Neuchatel, Terreaux 14, 2000 Neuchatel, Switzerland.

2 Agricultural Research Council, Private Bag XI34, Queenswood 0121, South Africa.

2 School of Biological and Conservation Sciences, University of KwaZulu-Natal, Private Bag X01,

Scottsville, Pietermaritzburg, 3209, South Africa.

3 Zoological Museum, University of Kiel, Hegewischstr. 3, D-24105 Kiel, Germany.

3 Department of Fife Sciences, Natural History Museum, Cromwell Road, Fondon SW7 5BD, UK.

* Corresponding author: [email protected]

2 Email: [email protected]

3 Email: [email protected]

1 um:lsid:zoobank.org:author:21E7A776-F85D-4C28-8930-0DF18E3A7CE0

2 um:lsid:zoobank.org: author: 1EBE2282-7334-47BB-A874-921C4120D8B8

3 urn:lsid:zoobank.org:author:B99AE0ED-FA89-4DFE-A658-lC8DF37F9FAB

Abstract. A new species of Fidelia Friese, 1899 is described from southern Africa: Fidelia ( Fideliopsis)

whiteheadi Fitman & Kuhlmann sp. nov. Diagnostic characters are provided to distinguish this species

from others in the genus, particularly from the closely related F. hessei', an updated description for

Fidelia hessei is also given. The host plant preferences and seasonal activity of F. whiteheadi Fitman &

Kuhlmann sp. nov. and F. hessei are discussed. Finally, a revised, illustrated key to species of the genus

Fidelia is presented.

Keywords. Bees, Fidelia whiteheadi sp. nov., South Africa, Namibia, pollinator.

Litman J.R., Eardley C.D. & Kuhlmann M. 2016. A new species of Fidelia Friese, 1899 (Hymenoptera,

Megachilidae), with a key to the species of the genus. European Journal of Taxonomy 174: 1-18. http://dx.doi.

org/10.5852/eit.2016.174

Introduction

The twelve members of the genus Fidelia Friese, 1899 are solitary, ground-nesting bees. A single species

is known from the xeric regions of the Atlas Mountains in Morocco (Warncke 1980; Whitehead &

Eardley 2003). The remaining eleven species are distributed throughout the winter and summer rainfall

areas of western and northwestern South Africa, as well as the deserts of Namibia; some observations

have also been made in Botswana.

European Journal of Taxonomy 174 : 1-18 ( 2016 )

Members of Fidelia are robust bees, ranging in size from approximately 7.5 to 20.0 mm (Whitehead &

Eardley 2003). Their pilosity may be white through yellow to orange, while the integument is mostly

brown to black, although also yellow on the clypeus, mandibles, labrum, and legs of some species. All

females and some males exhibit a pygidial plate, which is typically black but yellow in some species.

The male seventh tergum has either a simple or bifid apex bordered by lateral spines. Females have

a metasomal scopa, as well as a brush of long dense hairs on their hind basitarsi. The female hind

basitarsus itself is flattened and paddle-like and is used in nest excavation (Rozen 1977).

The genus Fidelia was originally described by Friese in an 1899 monograph (Friese 1899), while the

closely related genus Parafidelia Brauns, 1926 was later described by Brauns (1926). Michener (2000)

synonymized Fidelia and Parafidelia and proposed Parafidelia as a subgenus of Fidelia. He also described

a new subgenus, Fidelia ( Fideliana ) Michener, 2000, thus dividing Fidelia into three subgenera ( Fidelia ,

Fideliana and Parafidelia). Engel (2002) later removed two species from Fidelia (Parafidelia) and

placed them in a new genus, Fideliopsis Engel, 2002; Whitehead & Eardley (2003), however, considered

Fideliopsis as a subgenus of Fidelia , as did Michener (2007). The genus Fidelia is thus currently divided

into four subgenera, Fidelia (three species), Fideliana (two species), Parafidelia (two species) and

Fideliopsis (five species) (Michener 2007).

Extensive host plant records for the genus Fidelia were reported in Whitehead & Eardley (2003).

Although the records listed in Whitehead & Eardley (2003) do not distinguish between floral visits for

pollen and visits for nectar, the ensemble of these records suggest that most fideliines are oligolectic:

Fidelia fasciata Whitehead & Eardley, 2003 and F. hessei Whitehead & Eardley, 2003 on various species

of Grielum and Neuradopsis sp. (Neuradaceae); F. major Friese, 1911 on Grielum humifusum Thunb. and

Grielum sp.; F. braunsianaTriQSQ, 1905 on Berkheya spp. (Asteraceae); F. pallidula (Cockerell, 1935)

on Sisyndite spartea E.Mey. ex Sond. and Sisyndite sp. (Zygophyllaceae); F. ulrikei Wamcke, 1980

on Convolvulus trabutianus Schweinf. & Muschl. (Convolvulaceae); and F. villosa Brauns, 1902 on

various genera of Aizoaceae. F. kobrowi Brauns, 1905 and F. paradoxa Friese, 1899 have been collected

on Aizoaceae and Asteraceae but most records are on Aizoaceae; these bees are either oligolectic on

Aizoaceae or polylectic with a strong preference for Aizoaceae. Fidelia ornata (Cockerell, 1932) is

likely oligolectic on Aizoaceae (most visitation records are for Tribulocarpus sp.) (Whitehead & Eardley

2003), although other hosts are possible: Whitehead & Eardley (2003) list a single visitation record

for Tribulus sp. (Zygophyllaceae). While nearly all floral records for F. friesei (Brauns, 1926) are for

Sesamum triphyllum Welw. exAsch. and Sesamum sp. (Pedaliaceae), records for Salvia sp. (Lamiaceae)

and Crotalaria sp. (Fabaceae) have also been reported (Whitehead & Eardley 2003).

Members of Fidelia exhibit an unusual combination of morphological characters, including three

submarginal wing cells and a metasomal scopa. They also exhibit peculiar nesting behavior: they build

unlined nests, a behavior rarely seen in bees. A recently published molecular phylogeny identified the

genus Fidelia as one of the earliest branching lineages of the family Megachilidae (Litman et al. 2011).

Members of the genus Fidelia are thus phylogenetically significant bees and the discovery of a new

species is an important finding that justifies the description of a single new species in the context of a

recent revision of the genus (Whitehead & Eardley 2003).

Collecting trips undertaken by J.R.L. and C.D.E. to the Northern Cape Province in October 2008 and

February 2009 yielded two series of specimens that both key out to Fidelia ( Fideliopsis ) hessei in

Whitehead & Eardley’s (2003) key. Upon close inspection, however, both series of specimens exhibit

distinctly different features. A thorough re-examination of the type material used in the description of

F. hessei Whitehead & Eardley, 2003 indeed revealed two distinct but apparently closely related species.

We describe the new species here as Fidelia (Fideliopsis) whiteheadi Litman & Kuhlmann sp. nov.,

redescribe its close relative F. (F) hessei and provide a revised key to the species of Fidelia.

LITMAN J.R. et al. , New species of Fidelia from southern Africa

Materials and methods

The majority of the type material used in this description is deposited in the Iziko South African Museum

in Cape Town, South Africa; a limited number of specimens are also deposited in the Natural History

Museum of Neuchatel (Neuchatel, Switzerland), the Oberosterreiches Tandesmuseum, Biologiezentrum

(Tinz, Austria) and in the Titman-Praz Collection (Neuchatel, Switzerland). Morphological terminology

is taken from Michener (2007).

Abbreviations

We use the following abbreviations:

Bl = Body length (measured between anterior margin of clypeus and apex of metasoma)

Iw = Intertegular width (width of body measured between wing bases)

T = Metasomal tergum (when followed by a number, refers to number of tergum; for example, T1

refers to the first metasomal tergum)

S = Metasomal sternum (when followed by a number, refers to number of sternum; for example, S1

refers to the first metasomal sternum)

The following abbreviations are used for collections and institutions:

SAM = Iziko South African Museum, Cape Town, South Africa

MHNN = Natural History Museum of Neuchatel, Neuchatel, Switzerland

OOLM = Oberosterreiches Tandesmuseum, Biologiezentrum, Tinz, Austria

TPC = Titman-Praz Collection, Neuchatel, Switzerland

CSCF = Centre Suisse de Cartographie de la Faune, Neuchatel, Switzerland

The following abbreviations are used in the list of material examined:

NCP = Northern Cape Province, South Africa

WCP = Western Cape Province, South Africa

Photographs were taken with a Dino-Tite AM413T digital microscope at the Natural History Museum

of Tondon, as well as with a VHX-1000 Keyence digital microscope co-hosted by the MHNN and the

CSCF.

Results

Class Hexapoda Blainville, 1816

Order Hymenoptera Tinnaeus, 1758

Superfamily Apoidea Tatreille, 1802

Epifamily Anthophila Tatreille, 1804

Family Megachilidae Tatreille, 1802

Genus Fidelia Friese, 1899

Subgenus Fideliopsis Engel, 2002

Fidelia {Fideliopsis) whiteheadi Titman & Kuhlmann sp. nov.

urn:lsid:zoobank.org:act:B88E995E-lB96-4DD5-BC76-9E819BE92A4B

Figs 1A, C, E, 2A, C, 3A, C, E

Diagnosis

Fidelia (. Fideliopsis ) whiteheadi Litman & Kuhlmann sp. nov. can be distinguished from members

of the subgenera Fidelia and Fideliana by the length of the marginal cell of its forewing, which is

approximately 0.75x the distance between the apex of the marginal cell and the distal edge of the

wing (length of marginal cell less than half this distance in the subgenera Fidelia and Fideliana ).

European Journal of Taxonomy 174 : 1-18 ( 2016 )

F. whiteheadi Litman & Ku hlm a nn sp. nov. can be further distinguished from the two members of the

subgenus Parafidelia. It differs from Fidelia ( Parafidelia) friesei by the gentle curve of the female

mid-tibial spur (strongly sickle-shaped in female F. friesei) and by the unswollen forefemur and

forebasitarsus of the male (swollen in F. friesei) and from F. pallidula by the presence of two equal¬

sized mandibular teeth in females (apical and preapical tooth present in female F. pallidula) and by

the bifid, triangular T7 of the male (T7 long, parallel-sided and rounded at the apex in F. pallidula).

From other members of the subgenus Fidelia (Fideliopsis) (except F. hessei), F. whiteheadi Litman &

Kuhlmann sp. nov. can be distinguished by the unmodified forebasitarsus of the male (modified in

other members of F. (Fideliopsis)), the yellow, convex-sided pygidial plate of the female (black or

brown and concave-sided in other members of F. (Fideliopsis)) and the shape of the male S7 and S8

(see below for details).

Fidelia whiteheadi Litman & Kuhlmann sp. nov. is morphologically most similar to F. hessei.

F. whiteheadi Litman & Kuhlmann sp. nov. is smaller and less broad than F. hessei. Both females

and males of Fidelia whiteheadi Litman & Kuhlmann sp. nov. exhibit relatively short pilosity on the

thorax, metasoma and legs (Fig. 1A). In F. hessei , pilosity is comparatively longer (Fig. IB). In female

F. whiteheadi Litman & Kuhlmann sp. nov., hairs on the forebasitarsus are brushy and the tips of hairs

are often bent at a sharp angle (Fig. 1C); in comparison, the hairs on the forebasitarsus of F. hessei are

shaggy and unmodified (Fig. ID). The posterior margin of the pygidial plate in F. whiteheadi Litman &

Kuhlmann sp. nov. is evenly rounded and nearly triangular (Fig. IE); in F. hessei , it is slightly flattened

apically (Fig. IF).

A triangular protuberance at the base of the male T7 in F. whiteheadi Litman & Kuhlmann sp. nov. is

strongly elevated; when examined in profile, the posterior margin of the protuberance meets the surface

of the tergum at a nearly 90 degree angle (Fig. 2A). In F. hessei , the protuberance at the base of T7 is

less pronounced and the angle where the protuberance meets the surface of T7 is less sharp (Fig. 2B).

A half-moon shaped protuberance on the male S2 is present in F. whiteheadi Litman & Kuhlmann sp.

nov. but is only slightly elevated and often completely hidden by the pilosity of S2; the length of the

protuberance is about a quarter of the length of the sternum (Fig. 2C). In F. hessei , the half-moon shaped

protuberance is more strongly elevated and is usually easily visible despite the surrounding pilosity; the

length of the protuberance is about one-third of the length of the sternum (Fig. 2D).

The most striking differences between F. whiteheadi Litman & Kuhlmann sp. nov. and F. hessei are

found in the male hidden sterna and genitalia. In both F. whiteheadi Litman & Kuhlmann sp. nov. and

F. hessei , the male S7 bears two apicolateral processes but in F. whiteheadi Litman & Kuhlmann sp. nov.

the processes of S7 are relatively narrower, more weakly sclerotized, less hairy and shorter (Fig. 3A)

than those of F. hessei (Fig. 3B). The male S8 of F. whiteheadi Litman & Kuhlmann sp. nov. narrows

sharply in the posterior third (Fig. 3C), while that of F. hessei narrows more gradually (Fig. 3D). The

gonostylus of F. whiteheadi Litman & Kuhlmann sp. nov. is distinctly narrower and the basal outer edge

more rounded (Fig. 3E) than in F. hessei (Fig. 3F).

Etymology

This species is named for the late Vincent Booth Whitehead (1921-2005), whose extensive collection

of fideliine bees deposited at the Iziko South African Museum in Cape Town, South Africa provided the

majority of the material for this species description.

Type material

Holotype

NAMIBIA: Koes, Katzies, 09 Mar. 1982, V.B. Whitehead (SAM), SAM-HYM-B001841.

LITMAN J.R. et ah. New species of Fidelia from southern Africa

Paratypes (all specimens deposited in SAM unless otherwise noted)

NAMIBIA: 1 10 km N of Kalkrand, 24.03° S, 17.57° E, 07 Mar. 1980; 2 $$, 15 km S of Stampriet,

24.42° S, 18.48° E, 13 Apr. 1980; 1 $, 1 S, 16 km S of Stampriet, 24.42° S, 18.48° E, 8 Feb. 1984; 3 $ $,

1 S, 18 km S ofAroab, 26.93° S, 19.55° E, 11 Apr. 1980; 3 $$, 2 6V?, 25 km N of Koes, 25.75° S, 19.25° E,

27 Apr. 1981; 1 S, 25 km S of Stampriet, 24.25° S, 18.53° E, 13 Apr. 1980; 5 $$,30 km N of Koes,

25.72° S, 19.27° E, 12 Apr. 1980; 2 $$, 30 Ion S of Gochas, 25.05° S, 18.98° E, 12 Apr. 1980; 1 $, 1 8,

30km S ofLeonardville, 23.IT S, 18.87°E, 15Apr. 1980; 8 $$, 4 SS, 31 Ion S ofLeonardville, 23.77° S,

18.87° E, 15 Apr. 1981; 3 $$, 32 km S ofLeonardville, 23.77° S, 18.87° E, 18 Mar. 1979; 4 $$, 33 km E

of Gochas, 24.75° S, 19.12° E, 13 Apr. 1980; 3 $ $, 9 SS, 33 km NW of Vorstershoop, 25.63° S, 22.92° E,

29 Feb. 1980; 3 $$,34 km N of Koes, 25.7° S, 19.32° E, 12 Apr. 1980; 1 $, 4 km N ofLeonardville,

23.45° S, 18.77° E, 9 Feb. 1984; 1 $, 4 km N ofLeonardville, 23.45° S, 18.77° E, 10 Feb. 1984; 1 c$,5km

WofAus,26.65° S, 16.23°E, 6Oct. 1982;3 $$, 1 S, 5IonWofKlinghardtMts,27.38° S, 15.63°E,40ct.

1982; 2 $$, 5 km W of Leonardville, Farm 206,23.57° S, 18.32° E, 15 Oct. 1980; 6 $$, 51 km S ofAroab,

27.15° S, 19.48° E, 11 Apr. 1980; 5 $$, 3 S3, 6 km NW ofLeonardville, 23.47° S, 18.88° E, 15 Apr.

1981; 1 $, 61cm W ofLeonardville, 23.48° S, 18.73° E, 15 Apr. 1980; 1 $, 601cm S of Griinau, 28.15° S,

18.05° E, 17 Apr. 1980; 6 $$, 3 7 km NW ofLeonardville, 23.48° S, 18.72° E, 26 Apr. 1981; 3 $$,

Cucumis, 24.01° S, 18.42° E, 11 Feb. 1984; 1 $, 3 SS, Diamond Area No. 1, Klinghardt Mts, Spitzlcuppe

Sud, 26.5° S, 15.5° E, 19 Oct. 1974; 6 $ $, Diamond Area No. 1, Klinghardt Mts, Spitzlcuppe Sud, 26.5° S,

15.5° E, 21 Oct. 1974; 4 $$, 1 Grillental, 27.13° S, 15.38° E, 29 Sep. 1982; 6 $$, 11 Klinghardt

Mts, 27.38° S, 15.88° E, 1 Oct. 1982; 1 $, 2 SS, Klinghardt Mts, 27.38° S, 15.88° E, 2 Oct. 1982; 3 $ $,

2 c?c?, Koes, Katzies, 9 Mar. 1982; 1 S, Luderitz, Obib Dunes, 28.17° S, 16.8° E, 17 Sep. 1973; 1 $, Obib,

28.08° S, 16.75° E, 29 Oct. 1977; 1 $, 3 S3, Stampried 132, 24.38° S, 18.38° E, 8 Mar. 1982.

SOUTHAFRICA: 1 S, NCP, 11 km N of McCarthy’s Rust, 26.2° S, 22.57° E, 1 Mar. 1980; 4 $$, NCP,

14 1cm S of Kenhardt, 29.27° S, 21.04° E, 27 Nov. 1981; 1 $, NCP, 15 1cm N of Noenieput, 27.38° S,

20.1° E, 7 Feb. 1984; 2 $$, 1 g 9 NCP, 20 1cm NW of Kenhardt on Kakamas Rd, 29.14° S, 21.01° E,

27 Nov. 1981; 9 $$, NCP, 40 km N of Kakamas, Biesiepoort, 28.43° S, 20.6° E, 29 Apr. 1981; 1 $,NCP,

40 km SW of Kakamas, 29.05° S, 20.27° E, 6 Feb. 1984; 3 $ $, NCP, 40 km W of Van Zylsrus, 27.05° S,

21.67° E, 3 Mar. 1980; 1 $, NCP, 50 km S of Vioolsdrift, 29.17° S, 17.83° E, 20 Oct. 1980; 2 $$, NCP,

Breelcerie Dunes, Vanwyksvlei, 3 May 1985; 6 $$, 2 S3, NCP, Breekerie Dunes, Vanwyksvlei, 4 May

1985; 4 $$, NCP, Breekerie Dunes, Vanwyksvlei, 5 May 1985; 1 $, 3 S3, NCP, Jalckalsputs, 28.67° S,

16.95° E, 8 Oct. 1980; 1 3, NCP, Lelckersing, Richtersveld, 29° S, 17.1° E, 25 Sep. 1982; 4 $ $, 1 3, NCP,

Hotazel exit, Rt. 31,27°12.558' S, 22°57.264' E, 1 Feb. 2009 (2 $ $, 1 3 LPC; 2 $ $ MHNN); 3 $ $, NCP,

21cm NW of Hotazel, Rt. 31, 27°11.855' S, 22°55.909' E, 3 Feb. 2009 (1 $ LPC; 2 $$ OOLM).

Description

Female

Measurements. Bl= 10-11 mm ; Iw = 3 mm

Head. Broader than long. Integument black. Clypeus basally black, apically yellow, mandible yellow

with two red-brown teeth, upper tooth longer than lower. Head covered by dense white pilosity. Pilosity

near vertex either white or pale yellow. Clypeus convex, flattened medially. Clypeus densely, minutely,

superficially punctate apically; punctation gradually becoming larger and sparser basally, with spaces

between punctures reaching the diameter of one-half a puncture, rarely one puncture, at clypeal base.

Antennal scape and pedicel dark red-brown to black. Flagellum dorsally red-brown to pale yellow,

ventrally most often yellow, although occasionally also yellow-red.

Mesosoma. Integument dark red-brown to black. Mesosomal disc very densely punctate, almost no

spaces between punctures. Pilosity white to pale yellow, dense, generally short but often longer on

mesopleuron and scutellum. Wings colorless, venation light brown, papillate beyond veins. Integument

of legs red-brown, slightly lighter at apex of foretibia and forebasitarsus. Integument of hind femur

European Journal of Taxonomy 174 : 1-18 ( 2016 )

lighter in color than that of fore and mid femur. Foretrochanter, fore femur, most of foretibia covered in

white hairs. Apex of foretibia and forebasitarsus covered in shorter, golden hairs. Forebasitarsus with

brushy, erect hairs, often bent at a 90 degree angle; length of hairs at posterior base of basitarsus shorter

than width of basitarsus at its base (Fig. 1C). Mid and hind legs covered in long, dense white hairs.

Fig. 1 . Fidelia spp., A, C, E. Fidelia whiteheadi Litman & Kuhlmann sp. nov. (paratype). B, D,

F. F. hessei Whitehead & Eardley, 2003. A-B. Metasoma. C-D. Forebasitarsus. E-F. Pygidial plate.

Scale bars: 1 mm.

LITMAN J.R. et al ., New species of Fidelia from southern Africa

Hind basitarsus with long yellow-brown hairs on dorsal and ventral edges; dorsal hairs longer than

ventral hairs.

Metasoma. Integument red-brown to black on tergal discs, tergal margins transparent, light brown to

yellow. Pilosity white to yellow. Pilosity on T1 slightly longer than that of T2-T5. Pilosity on discs of

T2-T5 nearly uniform in density and length. Pilosity in longer tufts laterally on T1-T5. Pilosity longest

on T6, both on tergal disc and laterally. Lateral tufts on T6 seldom reach apex of pygidial plate (Fig. 1 A).

Outline of pygidial plate evenly rounded in dorsal view, nearly triangular in form. Color of pygidial plate

variable, from pale white-yellow to dark yellow (Fig. IE). Integument of sterna red-brown. Pilosity of

metasomal scopa white.

Male

Measurements. Bl = 9-11 mm; Iw = 2.5 mm

Same as for female except as follows:

Head. Clypeus uniformly dark red-brown. Pilosity on head unifo rml y white, even on vertex.

Fig. 2. Fidelia spp., SS- A, C. Fidelia whiteheadi Litman & Kuhlmann sp. nov. (paratype). B, D. F. hessei

Whitehead & Eardley, 2003. A-B. Protuberance at base of T7. C-D. Protuberance on S2, indicated by

red arrow. Scale bars: 1 mm.

European Journal of Taxonomy 174 : 1-18 ( 2016 )

Mesosoma. Pilosity on mesosoma white. Integument of forecoxa, foretrochanter and fore femur dark

brown, that of foretibia and foretarsus yellow. Pilosity on forelegs entirely white. Pilosity on forebasitarsus

approximately three times as long as width of basitarsus. On mid and hind legs, all segments dark brown

except apex of basitarsus and tarsal segments. Pilosity on mid and hind legs white.

Fig. 3. Fidelia spp., SS- A, C, E. Fidelia whiteheadi Litman & Kuhlmann sp. nov. (paratype). B, D,

F. F. hessei Whitehead & Eardley, 2003. A-B. S7. C-D. S8. E-F. Gonostyli. Scale bars: 1 mm.

LITMAN J.R. et al.. New species of Fidelia from southern Africa

Metasoma. Protuberance at base of T7 strongly elevated, making a 90 degree angle with surface of

tergum. Half-moon shaped protuberance on S2 weakly elevated, often hidden by pilosity on ventral

metasoma (Fig. 2C). S7 with two apicolateral processes; each process narrow, wealdy sclerotized and,

except at tip of each process, only sparsely hairy (Fig. 3A). Length of each process of S7 approximately

two-thirds as long as length of central disc of sternum (Fig. 3 A). S8 sharply narrowed in posterior third,

such that width of apex approximately one-fifth of width of base of sternum (Fig. 3C). S8 without

paraspiculae (Fig. 3C). Gonostylus narrow and, in dorsal view, basal outer edge rounded (Fig. 3E).

General distribution

Winter rainfall area of northwestern South Africa and southwestern Namibia; summer rainfall area of

north-central South Africa and southeastern and south-central Namibia (Fig. 4).

Floral hosts

Grielum grandillorum [sic, we interpret this as Grielum grandiflorum (L.) Druce], Grielum sinuatum,

Grielum sp., and Neuradopsis sp. (Whitehead & Eardley 2003).

Seasonal activity

Two periods of activity a year, one from February to May (in summer rainfall area) and the second from

September to November (in winter rainfall area) (Fig. 4).

Remarks

To clearly distinguish Fidelia (Fide hops is) whiteheadi Litman & Kuhlmann sp. nov. from Fidelia

(Fideliopsis) hessei, we also provide a description for the latter.

Fidelia ( Fideliopsis ) hessei Whitehead & Eardley, 2003

Figs IB, D, F, 2B, D, 3B, D, F

Holotype

SOUTH AFRICA: $, NCP, 29 km E of Port Nolloth, 29.18° S, 17.09° E, 18 Sep. 1983 (SAM) (as

designated by Whitehead & Eardley 2003), SAM-HYM-B001836.

Other material examined (all specimens deposited in SAM unless otherwise noted)

NAMIBIA: 2 Klinghardt Mts, 27.38° S, 15.88° E, 19 Oct. 1974.

SOUTH AFRICA: 2 $$, 5 Ion S of Ratelfontein, 32.13° S, 18.63° E, 25 Oct. 1978; 3 ??, 1 S, NCP,

27 km E of Port Nolloth, 29.3° S, 17.15° E, 18Sep. 1983; 5 $$, NCP, 27 Ion E of Port Nolloth, 29.283° S,

17.133° E, 25 Sep. 1982; 1 $, NCP, 30 Ion E of Port Nolloth, 29.3° S, 17.18° E, 2 Oct. 1981; 9

NCP, 36 miles Grass Flats, 29.38° S, 17.38° E, 04 Oct. 1980; 2 $?, 1 $, NCP, 38 Ion N of Concordia,

29.27° S, 18.0° E, 20 Nov. 1980; 1 $, NCP, Brandkaros, 28.47° S, 16.68° E, 29 Sep. 1982; 1 $, NCP,

Grasvlakte, 29.18° S, 17.42° E, 18 Sep. 1983; 3 $ §, NCP, Oograbies, 20 km E of Port Nolloth, 29.23° S,

17.13° E, 18 Nov. 1980; 11 ??, 3 $$, NCP, Wallekraal, Namaqualand, 30.38° S, 17.52° E, 1 Oct. 1950;

4 ??, 2 WCP, 10 km N of Graafwater, 32.07° S, 18.6° E, 21 Oct. 1981; 1 ?, WCP, 10 km N of

Leipoldtville, 32.13° S, 18.52° E, 25 Oct. 1978; 1 ?, WCP, 5 km E of Elands Bay, 32.28° S, 18.35° E, 08

Nov. 1978; 3 ??, 1 $, WCP, 7 km N of Leipoldtville, 32.18° S, 18.5° E, 08 Nov. 1978; 1 $, WCP, Elands

Bay, Baboon Point, 32.32° S, 18.32° E, 13 Nov. 1979; 1 <$, WCP, Graafwater, 7 km N of Leipoldtville,

32.18° S, 18.5° E, 08 Nov. 1978; 1 $, WCP, Paleisheuwel, 32.39° S, 18.43° E, 01 Nov. 2001; 1 $, WCP,

Paleisheuwel, 32.47° S, 18.72° E, 10 Nov. 1978; 3 ??, 4 S3, WCP, Paleisheuwel, 32.47° S, 18.72° E, 17

Nov. 1978; 3 $ ?, WCP, Skurfkop station, 31.88° S, 18.63° E, 21 Oct. 1981; 5 ? ?, 2 WCP, Skurflcop

station, 31.88° S, 18.63° E, 25 Oct. 1978; 2 $$, WCP, Skurfkop, 31.95° S, 18.62° E, 10 Nov. 1978; 1 ?,

NCP, 42 km S of Eeksteenfontein, 29°06.869' S, 17°25.694' E, 09 Oct. 2008 (LPC).

European Journal of Taxonomy 174 : 1-18 ( 2016 )

Note

The SAM database lists the date of collection for five male Fidelia hessei as “1982/04/25”. A close look

at the handwritten label on the specimens strongly suggests, however, that the labels do not read “25.

iv. 1982” but rather “25.ix. 1982”. Given the difficulty in reading the labels on these specimens, and given

the total absence of other specimens of F. hessei during the fall months, we consider these specimens to

have been collected in September and not April.

Description

Female

Measurements. Bl = 11-12 mm; Iw = 4 mm

Head. Broader than long. Integument on head and base of clypeus black. Clypeus apically dark to

yellow brown. Clypeus minutely, superficially punctate from base to apex. Mandible yellow with two

red teeth, upper tooth longer than lower. Head covered in dense white pilosity. Clypeus convex, flattened

medially. Antennal scape and pedicel dark red-brown. Dorsally, antennal flagellum dark red-brown

basally, becoming yellow-brown apically. Ventrally, flagellum yellow.

Mesosoma. Integument black. Mesosomal disc densely, minutely punctate. Pilosity dense, shaggy,

white to pale yellow, sometimes longer on mesopleuron and scutellum. Wings colorless to pale yellow,

venation light brown, papillate beyond veins.

Legs. Dark red-brown basally, becoming paler yellow-brown apically. Forecoxa, trochanter and femur

covered in white pilosity. Foretibia and foretarsus covered in golden pilosity. Pilosity on forebasitarsus

long, shaggy - length of hairs at posterior base of basitarsus approximately twice as long as width of

basitarsus at its base (Fig. ID). Mid and hind coxa, trochanter and femur covered in short white pilosity.

Midtibia, midtarsus and hind tibia covered in longer, denser white to white-yellow pilosity. Hind basitarsus

with long yellow-brown hairs on dorsal and ventral edges; dorsal hairs longer than ventral.

Metasoma. Integument black on tergal discs, tergal margins transparent, light brown to yellow. Pilosity

shaggy white to yellow-white, with pilosity on tergal discs often erect, while pilosity on tergal margins

lies flat (Fig. IB). Longer tufts of hair present laterally on T1-T6, with pilosity on T6 longer, with lateral

tufts almost reaching apex of pygidial plate. Outline of pygidial plate triangular, with flattened apex in

dorsal view (Fig. IF). Pygidial plate variable in color, from dark yellow to brown. Integument of sterna

red-brown. Hairs on metosomal scopa white.

Male

Measurements. Bl = 10-12 mm ; Iw = 2.5M mm

Same as for female except as follows:

Head. Integument on head and clypeus entirely dark brown to black.

Mesosoma. Pilosity white.

Legs. Forecoxa, trochanter and femur dark brown. Foretibia and foretarsus yellow. Pilosity on forelegs entirely

white. Pilosity on forebasitarsus long at base, approximately four times as long as width of forebasitarsus at its

base. Mid and hind legs and all segments dark brown except apex of basitarsus and tarsal segments. Pilosity

on mid and hind legs white, except on hind basitarsus, where ventral fringe of hairs is yellow white.

Metasoma. T7 dark brown to black. Protuberance at base of T7 usually weakly elevated, making a shallow

angle with respect to surface of tergum. Half-moon shaped protuberance on S2 strongly elevated, usually

LITMAN J.R. et al. , New species of Fidelia from southern Africa

visible through pilosity on ventral metasoma. S7 with two apicolateral processes (Fig. 3B); each process

broader, more strongly sclerotized, and more densely hairy than in F. whiteheadi Litman & Kuhlmann

sp. nov. Length of each process on S7 approximately one-half as long as length of central disc of sternum

(Fig. 3B). S8 narrows gradually in posterior third, such that width of apex is approximately one-third of

width of base of sternum (Fig. 3D). S8 without paraspiculae (Fig. 3D). Gonostylus distinctly broader and

basal outer edge more sharply angled (Fig. 3F) than in F. whiteheadi Litman & Kuhlmann sp. nov.

■4—'

* 1 —

H—

CtJ

■*->

(tl

Namibia

Windhoek

Botswana

Gobabis

Rehoboth

Gaborone

Marien'

Kgalagadi

Transfrontier

Park

Pretoria

Mahikeng

Johannesbui

Keetmanshoop

Welkom

Upington

Kimberley

Bloemfontein

Botshabelo

Springbok

Lesoth

South Africa

East London

Langebaan

Oudtshoorn

Port Elizabeth

Plettenberg

Bay

Knysnao o

6Jeffreys Bay

Hermanns

Mth,

Cape Town

Fidelia hessei

Swakopmund

fU ^

Namib-Naukluft

National Park

Solitaire

Lud 0 e, 0 (j

< Fm

Spengebiet

O Fidelia whiteheadi , autumn activity

# Fidelia whiteheadi, spring activity

Fig. 4. Distribution map of Fidelia hessei Whitehead & Eardley, 2003 (red dots), Fidelia whiteheadi

Litman & Kuhlmann sp. nov., autumn period of activity (light blue dots) and Fidelia whiteheadi

Litman & Kuhlmann sp. nov., spring period of activity (dark blue dots). The northernmost occurrence

of F. hessei shown on the map corresponds to morphologically ambiguous specimens collected in the

region around the Klinghardt Mountains of Namibia; the identification of the specimens from this region

should be corroborated with DNA data.

European Journal of Taxonomy 174 : 1-18 ( 2016 )

General distribution

Winter rainfall area of western and northwestern South Africa (Fig. 4).

Floral hosts

Grielum grandillorum [sic; we interpret this as Grielum grandiflorum\, Grielum sinuatum, Grielum sp.,

and Neuradopsis sp. (Whitehead & Eardley 2003).

Seasonal activity

One period of activity per year, from September to November.

Key to species 0/* Fidelia

(modified from Whitehead & Eardley 2003)

Note: in couplets where color of vestiture is mentioned, color refers to that of freshly eclosed specimens.

In older specimens, color of vestiture is often faded compared to that of younger specimens.

1. Marginal cell of forewing shorter than half distance between apex of marginal cell and distal wing

edge; female pygidial plate yellow and covering entire dorsal surface of T6; southern Africa and

Morocco (. Fidelia sensu stricto and Fide liana ).2

- Marginal cell of forewing shorter (0.75x) than distance between marginal cell and distal end of

forewing; female pygidial plate yellow or black, large but not covering entire dorsal surface of T6, a

little vestiture visible on each side of pygidial plate, except F. hessei which resembles Fidelia sensu

stricto; southern Africa ( Parafidelia and Fideliopsis) .6

2. Female clypeus black to mostly black; male with foretibia and forebasitarsus unmodified ( Fideliana )

- Female clypeus yellow to mostly yellow; male forebasitarsus with distinct groove, foretibia expanded

(. Fidelia sensu stricto ).4

3. Female vestiture mostly orangish and black, T5 black; male mandible with an apical and a sub-

apical tooth. F. braunsiana Friese, 1905

- Female vestiture, including T5, mostly white; male mandible with a single tooth.

. F. ulrikei Warncke, 1980

4. Female T5 with orange vestiture; male T7 as in Fig. 5A; male forebasitarsus weakly expanded

(Fig. 5B), distinctly less than half as wide as maximum length. F. kobrowi Brauns, 1905

- Vestiture on female T5 white to pale yellow; male forebasitarsus greatly expanded, more than half

as wide as maximum length.5

5. Distal part of female pygidial plate black; scopa yellowish; male with white vestiture; T7 as in

Fig. 5C; male forebasitarsus expanded posteriorly (anterior and posterior edges more or less parallel),

as in Fig. 5D. F. villosa Brauns, 1902

- Female pygidial plate completely yellow; scopa white; male with white to yellow vestiture; T7 as in

Fig. 5E; male forebasitarsus expanded more strongly posteriorly (posterior edge strongly convex),

as in Fig. 5F. F. paradoxa Friese, 1899

6. Female middle tibial spur strongly sickle shaped; hind tibia with black pubescence; male T7 as

in Fig. 6 A; male fore femur strongly swollen (length double width) and forebasitarsus moderately

swollen (0.4x as long as wide). F. friesei (Brauns, 1926)

LITMAN J.R. et al.. New species of Fidelia from southern Africa

- Female middle tibial spur gently curved; hind tibia with pallid pubescence; male forefemur

unmodified (width distinctly less than half length); forebasitarsus unmodified or strongly swollen

(0.6-0.7x as long as wide).7

Fig. 5. Fidelia spp., 88- A-B. Fidelia kobrowi Brauns, 1905. A. T7. B. Forebasitarsus. C-D. F. villosa

authorship?. C. T7. D. Forebasitarsus. E-F. F. paradoxa Friese, 1899. E. T7. F. Forebasitarsus. Scale

bars: 1 mm

European Journal of Taxonomy 174: 1-18 (2016)

7. Female mandible with an apical and a preapical tooth; clypeus and labrum black; male T7 as in

Fig. 6B; forebasitarsus unmodified. F. pallidula (Cockerell, 1935)

- Female mandible with two large, equal-sized teeth; clypeus yellow and labrum at least partly yellow

(only ventral half yellow in F. borearipa Whitehead & Eardley, 2003); male T7 as in Figs 6C-E, 7A,

C; forebasitarsus enlarged, except unmodified in F. hessei and moderately enlarged in F. borearipa

(Fideliopsis) .8

Fig. 6. Fidelia spp., S3- A-E. T7. A. Fidelia friesei (Brauns, 1926). B. F. pallidula (Cockerell, 1935).

C. F. hessei Whitehead & Eardley, 2003. D. F. whiteheadi Litman & Kuhlmann sp. nov. E. F. fasciata

Whitehead & Eardley, 2003 (paratype). F. F. fasciata , forebasitarsus (paratype). Scale bars: 1 mm.

LITMAN J.R. et ah. New species of Fidelia from southern Africa

8. Female pygidial plate yellow and convex laterally, with no T6 vestiture visible on side of pygidial

plate in dorsal view; male foretarsus not modified, T7 as in Fig. 6C or 6D.9

- Female pygidial plate black or brown and concave laterally, with a little T6 vestiture visible on side

of pygidial plate in dorsal view; male foretarsus modified.10

9. Female pygidial plate with flattened apex (Fig. IF); pilosity on female forebasitarsus long, shaggy

(Fig. ID); length of apicolateral process of male S7 approximately one-half as long as length of

central disc of sternum (Fig. 3B); male S8 narrows gradually in posterior third (Fig. 3D).

. F. hessei Whitehead & Eardley, 2003

- Female pygidial plate nearly triangular in form (Fig. IE); pilosity on female forebasitarsus brushy,

erect (Fig. 1C); length of apicolateral process on male S7 approximately two-thirds as long as length

of central disc of sternum (Fig. 3A); male S8 narrows sharply in posterior third (Fig. 3C).

. F. whiteheadi Litman & Kuhlmann sp. nov.

10. Vestiture on metasomal T2-T5 in female and T3-T6 in male black anteriorly, white posteriorly;

female hind tibia white dorsally; male T7 as in Fig. 6E; male forebasitarsus as in Fig. 6F.

. F. fasciata Whitehead & Eardley, 2003

- Vestiture on female metasoma mostly yellowish, sometimes posterior region of T2-T4 white, and

male completely white; female hind tibia yellowish dorsally.11

Fig. 7. Fidelia spp., 33- A-B. Fidelia ornata (Cockerell, 1932). A. T7. B. Forebasitarsus. C-D. F. major

Friese, 1911. C. T7. D. Forebasitarsus. Scale bars: 1 mm.

European Journal of Taxonomy 174: 1-18 (2016)

11. Female hind basitarsus mostly black; T7 as in Fig. 7A; male forebasitarsus as in Fig. 7B.

. F. ornata (Cockerell, 1932)

- Female hind basitarsus orangish above (orange or black below); T7 not as depicted in Fig. 7A; male

forebasitarsus different.12

12. Female hind basitarsus orange below; T7 as in Fig. 7C; male forebasitarsus as in Fig. 7D.

. F. major Friese, 1911

- Female hind basitarsus black below; T7 resembles that in Fig. 6E; male forebasitarsus moderately

expanded (width 0.4x length). F. borearipa Whitehead & Eardley, 2003

Discussion

According to the criteria outlined by Engel (2004), Fidelia whiteheadi Litman & Kuhlmann sp. nov.

clearly belongs to the subgenus Fideliopsis : the mandible of both male and female are bifid, with the

teeth long, slender and divergent and the medioapical process of the male T7 shallowly emarginate at

the apex. Within Fideliopsis , F. whiteheadi Litman & Kuhlmann sp. nov. is most similar to F. hessei ;

both exhibit a yellow to pale yellow female pygidial plate, unmodified male protarsi, short apicolateral

processes of the male S7 that taper from a broad base and whose lengths do not surpass the width of the

central disc and an absence of paraspiculae on the male S8 (Whitehead & Eardley 2003; Engel 2004).

F. whiteheadi Litman & Kuhlmann sp. nov. can be distinguished from F. hessei , however, by its smaller

size, shorter pilosity (Fig. 1A-B), the nature of the pilosity on the female forebasitarsus (Fig. 1C-D), the

shape of the apex of the female pygidial plate (Fig. 1E-F), the elevation of the protuberance at the base

of the male T7 (Fig. 2A-B), the elevation and width of the protuberance on male S2 (Fig. 2C-D), and,

most notably, the shape of the male S7, S8 and gonostylus. (Fig. 3A-F).

Fidelia hessei appears to have a single period of activity each year, in the spring months (September-

November) of the winter rainfall areas of the Western and Northern Cape provinces of South Africa

(Fig. 4). Fidelia whiteheadi Litman & Kuhlmann sp. nov., on the other hand, appears to have two periods

of activity a year, one in the spring months (September-November) of the winter rainfall areas of the

Northern Cape Province and southwestern Namibia and the second in the autumn months (February-

May) of the summer rainfall areas of the Northern Cape Province and the south central and southeastern

regions of Namibia (Fig. 4). A few isolated specimens from spring-emerging F. whiteheadi Litman &

Kuhlmann sp. nov. were found in localities more typical of autumn-emerging F. whiteheadi Litman &

Kuhlmann sp. nov. (Fig. 4). The distribution of F. hessei and that of spring-emerging F. whiteheadi

Litman & Kuhlmann sp. nov. are sympatric through a portion of their distributions, in the Northern Cape

Province of South Africa near the Namibian border.

A handful of specimens exhibit morphological characters intermediate between the two species. Curiously,

these specimens are not found in the region of sympatry between the two species but rather in the region

of the Klinghardt Mountains, at the northern extreme of the distribution of spring-emerging F. whiteheadi

Litman & Kuhlmann sp. nov., where few, if any, F. hessei occur (Fig. 4). All examined females from

this region appear to be F. whiteheadi Litman & Kuhlmann sp. nov. Based on specimen size, overall

pilosity, the shape of the protuberance on S2, and the shape of the pygidial plate, males also appear to

be F. whiteheadi Litman & Kuhlmann sp. nov.; an examination of male S7 and S8, however, strongly

suggests F. hessei in some specimens. It is not clear whether these specimens are simply small, atypical

F. hessei or F. whiteheadi Litman & Kuhlmann sp. nov. with unusual hidden sterna. If these specimens

are indeed F. hessei , the geographic distribution of this species would extend well into the winter rainfall

area of southwestern Namibia (Fig. 4). While some specimens were identified as F. whiteheadi Litman &

Kuhlmann sp. nov. and others as F. hessei , these determinations are provisional; the use of genetic data

will be an important element for the identification of these specimens in the future.

LITMAN J.R. et al., New species of Fidelia from southern Africa

According to Whitehead & Eardley (2003), plant visitation records for F. whiteheadi Litman & Kuhlmann

sp. nov. and F. hessei include Grielum grandillorum [sic, we interpret this as Grielum grandiflorum],

Grielum sinuatum, Grielum sp. and Neuradopsis sp. In order to determine whether either F. whiteheadi

Litman & Kuhlmann sp. nov. or F. hessei demonstrate a particular preference for any of these hosts,

pollen slides were prepared using pollen taken from 24 female specimens representing both species.

Pollen identification proved more difficult than anticipated and these slides have yet to be fully analyzed.

They are deposited at the SAM and we encourage those interested to examine these slides for further

clues as to the host plant preferences of F. whiteheadi Litman & Kuhlmann sp. nov. and F. hessei.

Acknowledgements

Discovery of this species would not have been possible without grants from the Graduate School and

the Mario Einaudi Center at Cornell University to J.R.L. and a grant through the Griswold Endowment

(Department of Entomology, Cornell University) to J.R.L., as well as a grant from the National

Geographic Society (8448-08) to Bryan N. Danforth. We thank the Izilco South African Museum for the

generous loan of specimens. Finally, we thank two anonymous reviewers whose comments substantially

improved this manuscript.

References

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Engel M.S. 2002. Phylogeny of the bee tribe Fideliini (Hymenoptera: Megachilidae), with the description

of a new genus from southern Africa. African Entomology 10 (2): 305-313.

Engel M.S. 2004. Fideliine phylogeny and classification revisited (Hymenoptera: Megachilidae).

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of the mature larva of Parafidelia (Hymenoptera, Apoidea). American Museum Novitates 2637: 1-15.

Warncke K. 1980. Fidelia, eine far die Westpalaarktis neue Bienengattung. Mitteilungen der Miinchner

Entomologischen Gesellschaft 70: 89-94. Available from http://biodiversitvlibrarv.org/page/28992198

[accessed 19 Dec. 2015]

Whitehead V.B. & Eardley C.D. 2003. African Fideliini: Genus Fidelia Friese (Hymenoptera: Apoidea:

Megachilidae: Fideliinae). Journal of the Kansas Entomological Society 76: 250-276.

Manuscript received: 28 September 2015

Manuscript accepted: 28 October 2015

Published on: 10 February 2016

Topic editor: Koen Martens

Desk editor: Charlotte Thionois