European Journal of Taxonomy 34: 1-34

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2Q12 • Kuhlmann M.

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Research article

um:lsid:zoobank.org:pub:B4F3BFEB-6QAE-4F15-BBlD-41B3E7BEC299

Revision of the South African endemic bee genus Redivivoides

Michener, 1981 (Hymenoptera: Apoidea: Melittidae)

Michael KUHLMANN

Department of Fife Seienees, The Natural History Museum,

Cromwell Road, Eondon, SW7 5BD, United Kingdom.

E-mail: [email protected]

um: Isid: zoobank, org: author: B99 AEQED-FA89-4DFE-A65 8-1C 8DF 3 7F 9FAB

Abstract. The South Afriean endemie bee genus Redivivoides Miehener, 1981 is revised and redefined.

The genus eomprises seven speeies, six of whieh are deseribed here as new: Redivivoides capensis

sp. nov. $(?, R. eardleyi sp. nov. $, R. kamieskroonensis sp. nov. $, R. karooensis sp. nov.

R. namaquaensis sp. nov. andR. variabilis sp. nov. Akey to speeies is provided.

Key words. Redivivoides, bees. South Afriea, new speeies, taxonomy.

Kuhlmann M. 2012. Revision of the South African endemic bee genus Redivivoides Michener, 1981 (Hymenoptera:

Apoidea: Melittidae). European Journal of Taxonomy 34:1-34. http://dx.doi.org/10.5852/eit.2012.34

Introduction

The bee genus Redivivoides was deseribed by Miehener (1981) based on a single speeies, R. simulans

Miehener, 1981, eolleeted in the winter rainfall region of western South Afriea. Two other speeies were

only known from a few female speeimens and remained undeseribed. Sinee then mueh more material

of this genus has beeome available representing now a total of seven speeies, six of whieh are deseribed

here as new. The morphologieal diversity of the newly deseribed speeies required a re-evaluation of the

eharaeters defining Redivivoides as a genus.

Redivivoides belongs to the subfamily Melittinae Sehenek, 1860 and the tribe Melittini Sehenek, 1860

that also ineludes the genera Melitta Kirby, 1802 and Rediviva Friese, 1911 (Miehez et al. 2009).

Miehener (1981) suggested that Redivivoides is the sister-group of Rediviva and the elose relationship

of both genera was later eonfirmed by phylogenetie studies (Danforth et al. 2006; Miehez et al. 2009).

However, unlike Redivivoides, the females of Rediviva eolleet fioral oil that is mixed with pollen for nest

provisioning from a range of oil-produeing fiowers, with Diascia (Serophulariaeeae) as their prineipal

fioral host (Whitehead & Steiner 2001; Pauw 2006; Whitehead et al. 2008). In several Rediviva speeies

the forelegs are elongate, sometimes longer than the entire body, and the lengths of fioral spurs and

bee legs in some eases show eo-variation at the population level, suggesting eo-evolution (Steiner &

Whitehead 1990, 1991). Beeause Melittidae are the most basal bees (Danforth et al. 2006), Redivivoides

and Rediviva are a key group for understanding the evolution of oil-eolleeting in bees (Miehez et al.

2009) and they might also help to understand the origin of the unusual bee diversity in the Greater Cape

Floristie Region (Kuhlmann 2009).

European Journal of Taxonomy 34: 1-34 (2012)

The goal of this publication is to describe the six new species and to provide a key for species identification

to facilitate further research on this fascinating group of bees. Based on the descriptions and the analysis

of the morphology of the new species, Redivivoides is redefined as a genus and its relationship to its

putative sister group, the oil-collecting Rediviva bees, is briefiy discussed.

Material and methods

Terminology for the description of species is based on Michener (2007) for general morphology.

Puncture density is expressed as the relationship between puncture diameter (d) and the space between

them (i), such as i = 1.5 d or i < d. The following abbreviations were used for morphological structures:

T = metasomal tergum

S = metasomal sternum

Body length was measured from the vertex to the apex of the body.

Acronyms for collections (after Arnett et al. 1993) from which specimens were borrowed or deposited,

are as follows:

AMGS

CUIC

EMUS

LPCT

SAMC

SANG

RCMK

Albany Museum, Grahamstown, South Africa

Cornell University Collection, Ithaca, USA

Entomological Museum, Utah State University, USA

Eaurence Packer collection, Toronto, Canada

South African Museum, Cape Town, South Africa

South African National Collection of Insects, Pretoria, South Africa

research collection of Michael Kuhlmann, Eondon, UK

Nomenclature of plant names follows Germishuizen & Meyer (2003). If not given on the labels,

coordinates of collecting sites are given in square brackets and places were identified using Microsoft

Encarta® World Atlas (version 10, 2001). Geographical coordinates are given in the following format:

degree.minutes.seconds. Distribution maps were generated using DMAP V7.2 I www.dmap.co.uk l .

Scanning electron microscopy (SEM) was carried out using a Eeo 1455VP. The backscattered electron

images were taken under low vacuum (variable pressure) of uncoated specimens. In this mode electron-

poor organic material appears darker than electron-rich material (e.g. soil dust particles) consisting of

heavier chemical elements.

Results

Classis Hexapoda Blainville, 1816

Ordo Hymenoptera Einnaeus, 1758

Superfamilia Apoidea Eatreille, 1802

Epifamilia Anthophila Eatreille, 1804

Familia Melittidae Schenck, 1860

Subfamilia Melittinae Schenck, 1860

Tribus Melittini Schenck, 1860

Genus Redivivoides Michener, 1981

KUHLMANN M., Revision of the bee genus Redivivoides

RedivivoidesMiQhQWQx, 1981

Redivivoides Michener, 1981: 42-44, figs 80, 82-84 (p. 43), 97 (p. 48).

Type species: Redivivoides simulans Michener, 1981, by original designation.

The diagnosis of Redivivoides given by Michener (1981) was solely based on R. simulans, which was

the only known species of the genus at that time. However, in some respects R. simulans is an exception

within the genus so a redefinition of Redivivoides is required in the light of the six new species described

in this paper (see discussion).

A comprehensive phylogenetic analysis of Redivivoides and related genera is in preparation to investigate

the intra-generic relationships, so for convenience the species are here listed in alphabetical order.

Redivivoides capensis sp. nov.

Figs lE-F, 2, 3, 4

um:lsid:zoobank.org:act:60C46E16-6E85-43BE-AA18-A7977B6CFQ97

Diagnosis

Males and females of R. capensis sp. nov. can be separated from other Redivivoides species by their

extensively and brightly red metasomal terga (Figs 2D, 3C) in combination with an impunctate and

densely shagreened scutum (Figs 2C, 3B).

Fig. 1. — A-B. Macropis europaea Wamcke, 1973, $. A. Hind leg, dorsal view. B. Hind basitarsus,

dorsal view. — C-D. Rediviva intermixta (Cockerell, 1934), C- Hind leg, dorsal view. D. Hind

basitarsus, dorsal view. — E-F. Redivivoides capensis sp. nov., $. E. Hind tibia, dorsal view. F. Hind

basitarsus, dorsal view. — G-H. Melitta arrogans (Smith, 1879), $. G. Hind tibia, dorsal view. H. Hind

basitarsus, dorsal view. Scale bar: 1 mm.

European Journal of Taxonomy 34: 1-34 (2012)

Etymology

Named after the Cape of Good Hope (“the Cape”).

Type material (18 speeimens)

Holotype

M. Kuhlmaim leg. 15 Aug. 2010 / Redivivoides spee. 2, M. Kuhlmann det. 2011 (SAMC). SOUTH

AFRICA: Farm Papkuilsfontein, 20 km S Nieuwoudtville, Fynbos, 31°33’16” S, 19°08’31” E, 680 m.

Paratypes

SOUTH AFRICA: 4 $ $ same date as Holotype (RCMK); 1 $, N. Cape, Fynbos, 2 km W Nieuwoudtville,

near Grasberg [31°2U05” S, 19°05’49” E], 742 m, 23 Aug. 2003, K. Timmermann (RCMK); 1 $, N.

Cape, Nieuwoudtville, Flower Reserve East [31°2U56” S, 19°08’52” E], 735 m, 11 Sep. 2003, M.

Kuhlmann (RCMK); 1 $, 2 33, N. Cape, Nieuwoudtville, Glen Eyon, Renosterveld [31°24’03” S,

19°08’34” E], 700 m, 7 Aug. 2004, M. Kuhlmann (RCMK); 2 33 , N. Cape, Nieuwoudtville, Glen

Eyon, Renosterveld [31°24’03” S, 19°08’34” E], 700 m, 11 Aug. 2004, M. Kuhlmann (RCMK); 4 $

20 km S Nieuwoudtville, Farm Papkuilsfontein, Fynbos [31°33’16” S, 19°08’31” E], 680 m, 27 Aug.

2010, M. Kuhlmann (SANC, RCMK); 1 5, vanRhynsdorp. Cape [31°36’ S, 18°44’ E], 1929, Dr. Brauns

(SAMC); 1 ?, Clanwilliam, Ramskop [32°10’ S, 18°53’ E], 24 Jul. 1985, V.B. Whitehead (SAMC).

Fig. 2. Redivivoides capensis sp. nov., $. A. Eateral view. B. Head. C. Seutum and seutellum.

D. Metasoma (dorsal view). Seale bar: 1 mm.

KUHLMANN M., Revision of the bee genus Redivivoides

Fig. 3. Redivivoides capensis sp. nov., S- A. Lateral view. B. Seutum and seutellum. C. Metasoma

(dorsal view). D. S6 (ventral view). E. S7 (dorsal view). F. S8, apieal end (baekseattered eleetron

image). G. Genitalia (dorsal view). H. Genitalia (lateral view). Seale bar: 1 mm.

European Journal of Taxonomy 34: 1-34 (2012)

Description

Female

Body length. 9.0-10.5 mm .

Head. Head slightly wider than long. Integument blaek exeept tips of mandible partly dark reddish-

brown. Faee sparsely eovered with long, whitish-grey to brown, ereet hairs intermixed with blaek hairs

espeeially along the inner eye margins and on vertex (Fig. 2B). Clypeus mostly flat, apieally almost

impunetate; medially eovered with medium-sized punetures that beeome gradually smaller and denser

towards the lateral and upper margins; surfaee between punetures shiny (Fig. 2B). Malar area medially

narrow, almost linear. Antenna blaek, ventrally orange to reddish-brown.

Mesosoma. Integument blaek. Mesoseutum impunetate, finely shagreened and matt (Fig. 2C).

Mesoseutum, seutellum, metanotum, mesepistemum and propodeum eovered with long yellowish-

brown ereet hairs, on the dise of mesoseutum some blaek hairs intermixed.

Wings. Yellowish-brown; wing venation dark brown.

Legs. Integument blaek to dark reddish-brown. Vestiture of femora whitish-yellow, on tibiae and tarsi

dark brown to blaek, seopae dark brown, ventrally hairs white to greyish-yellow.

Metasoma. Integument blaek, exeept T1 to a variable extent apieally red, T2 red with a blaek spot

anterior-medially, T3 apieally with a red margin (Fig. 2D). T1 eompletely and T2 on dise sparsely

eovered with a few long ereet yellowish-white hairs; T3 - T4 eovered with short yellow to blaek hairs;

apieal tergal hair band missing on Tl, on T2 - T4 broadly white (Fig. 2A, D). Prepygidial fimbria

dorsally blaek, laterally partly white, pygidial fimbria blaek. Terga impunetate, finely shagreened with

a silky shine (Fig. 2D).

Male

Body eength. 8.5-10.0 mm.

Head. Head slightly wider than long. Integument blaek exeept tip of mandible partly dark reddish-

brown. Faee sparsely eovered with long, whitish-grey to brown, ereet hairs intermixed with blaek hairs

espeeially along the inner eye margins and on vertex. Clypeus mostly flat, apieally almost impunetate;

medially eovered with medium-sized punetures that beeome gradually smaller and denser towards the

lateral and upper margins; surfaee between punetures shiny. Malar area medially narrow, almost linear.

Antenna blaek, ventrally reddish-brown.

Mesosoma. Integument blaek. Mesoseutum impunetate, finely shagreened and matt (Fig. 3B).

Mesoseutum, seutellum, metanotum, mesepistemum and propodeum eovered with long white to

yellowish-grey ereet hairs, on the seutellum blaek hairs intermixed (Fig. 3A).

Wings. Slightly yellowish-brown; wing venation brown.

Legs. Integument blaek, tibiae and tarsi partly dark red-brown. Vestiture whitish.

Metasoma. Integument blaek, exeept Tl to a variable extend apieally red, T2 red with a blaek spot

anterior-medially, T3 either like T2 or in some speeimens only apieally with a red margin, T4 in some

males with red basal and apieal margins (Fig. 3C). Tl eompletely and T2 - T3 on dise sparsely eovered

with a few long ereet yellowish-white hairs; T4 - T6 eovered with short blaek hairs; apieal tergal hair

band missing on Tl, on T2 -T5 narrow and sparsely white (Fig. 3C). Terga impunetate, finely shagreened

with a silky shine (Fig. 3C).

KUHLMANN M., Revision of the bee genus Redivivoides

Terminalia. Genitalia (Fig. 3G-H), S6 (Fig. 3D), S7 (Fig. 3E) and terminal plate of S8 (Fig. 3F) as

illustrated.

Distribution

This species is only known from a few localities from Clanwilliam to the Nieuwoudtville area (Fig. 4).

Floral hosts

All specimens collected on the farms Papkuilsfontein and Glen Lyon were collected on Polygalaceae:

Nylandtia scoparia.

Seasonal activity

July - September.

20E 25E 30E

Fig. 4. Distribution of Redivivoides capensis sp. nov. andR. karooensis sp. nov. in South Africa.

European Journal of Taxonomy 34: 1-34 (2012)

Redivivoides eardleyi sp. nov.

Figs 5, 9

um:lsid:zoobank.org:act:543F31E2-5A26-4D78-AQ7A-674CA7997F19

Diagnosis

Females of R. eardleyi sp. nov. ean be separated from other Redivivoides speeies by a eombination of

the following eharaeters: metasomal terga blaek to brown, T2 sparsely punetate with minute punetures

and surfaee between punetures smooth and shiny, white apieal tergal hair bands present (Fig. 5D),

prepygidial and pygidial fimbria dark brown to blaek (Fig. 5D). The male is unknown.

Etymology

Named after Connal D. Eardley, Pretoria, who eolleeted this speeies and to honour his outstanding

eontribution to Afriean bee taxonomy.

Type material (3 speeimens)

Holotype

?, 13 Sep. 2011, C. Eardley (SANG). SOUTH AFRICA: N. Cape, 6 km E Pofadder, 29°06’26” S,

19°27’13”E.

Fig. 5. Redivivoides eardleyi sp. nov., $. A. Eateral view. B. Head. C. Seutum and seutellum.

D. Metasoma (dorsal view). Seale bar: 1 mm.

KUHLMANN M., Revision of the bee genus Redivivoides

Paratypes

SOUTH AFRICA: 1 same date as holotype (RCMK); 1 N. Cape, 21.6 km E Springbok [29°34’ S,

18°04’ E], 1006 m, 13 Sep. 2011, E. Packer (EPCT).

Description

Female

Body LENGTH. 11.0 mm.

Head. Head wider than long. Integument black. Face sparsely covered with long, whitish-grey, erect

hairs, along the inner eye margins and on vertex intermixed with black hairs (Fig. 5B). Clypeus convex

in profile, apically impunctate; medially covered with fine punctures that become gradually smaller and

denser towards the clypeal margins; surface between punctures smooth and shiny (Fig. 5B). Malar area

medially narrow, a lm ost linear. Antenna black.

Mesosoma. Integument black. Mesoscutal disc between punctures smooth and shiny; disc densely

(i = 0.5-1.0 d) and finely punctate (Fig. 5C). Mesoscutum, scutellum, metanotum, mesepistemum and

propodeum covered with long yellowish-white erect hairs, on the disc of mesoscutum intermixed with

black hairs.

Wings. Yellowish-brown; wing venation reddish brown.

Eegs. Integument black. Vestiture whitish to brown, scopae yellowish to dark brown.

Metasoma. Integument black, apical margins of T2 - T4 narrowly reddish-brown (Fig. 5D). T1 with a

few long erect whitish hairs; discs of T2 - T4 sparsely covered with very short erect white to dark brown

hairs; apical tergal hair bands on T1 - T4 broad and white; prepygidial and pygidial fimbriae black (Fig.

5A, D). T1 almost impunctate, polished and shiny, T2 - T4 shiny, with very fine, superficial and sparse

punctation that becomes progressively denser on apical tergae (Fig. 5D).

Male

Unknown.

Distribution

There is only a single record of this species from the Karoo in the summer rainfall area (Fig. 9).

Floral hosts

Unknown.

Seasonal activity

September.

Redivivoides kamieskroonensis sp. nov.

Figs 6, 9

um:lsid:zoobank.org:act:C65BQCFE-D36A-4C57-BE42-7QQ9CEQF357D

Diagnosis

Females of R. kamieskroonensis sp. nov. can be separated from other Redivivoides species by their

extensively and brightly red metasomal terga (Fig. 6D) in combination with the head that is distinctly

wider than long (Fig. 6B). Unlike other species the propodeal triangle is weakly shagreened. The male

is unknown.

European Journal of Taxonomy 34: 1-34 (2012)

Etymology

Named after the town Kamieskroon in NW South Afriea, in the vieinity of whieh this speeies was

diseovered.

Type material (2 speeimens)

Holotype

16-17 Sep. 2001, C. EmdXQy!Redivivoides sp. 3, det. C.D. Eardley 2008/(bluelabel): SANC-Pretoria,

Database No: HYMA 08832. SOUTH AFRICA: N Cape, Dassiefontein Farm, 14 km E Kamieskroon,

30°09’ S, 17°59’E.

Paratype

SOUTH AFRICA: 1 $, NCP, Dassiefontein Farm, 14 road km E Kamieskroon [30°09’31” S,

17°59’96” E], 760 m, 16-17 Sep. 2001 (CUIC).

Description

Female

Body length. 12.0 mm.

Fig. 6. Redivivoides kamieskroonensis sp. nov., $. A. Fateral view. B. Head. C. Seutum and seutellum.

D. Metasoma (dorsal view). Seale bar: 1 mm.

KUHLMANN M., Revision of the bee genus Redivivoides

Head. Head much wider than long. Integument black except tips of mandible partly dark reddish-

brown. Face sparsely covered with long, whitish-grey to brown, erect hairs intermixed with black hairs

(Fig. 6B). Clypeus almost flat, slightly convex in profile in the lower part and slightly concave in the

upper part, apico-medially and apical margin impunctate; clypeus otherwise densely (i < d) covered

with small punctures that become gradually smaller and denser towards the clypeal margins; surface

between punctures shiny (Fig. 6B). Malar area medially about 1/4 to 1/5 as long as width of mandible

base. Antenna black, ventrally reddish-brown.

Mesosoma. Integument black. Mesoscutal disc between punctures smooth and shiny; disc densely

(i = 0.5-1.0 d) and finely punctate (Fig. 6C). Mesoscutum, scutellum, metanotum, mesepistemum

and propodeum covered with long yellowish to whitish-grey erect hairs, on the disc of mesoscutum

intermixed with black hairs.

Wings. Yellowish-brown; wing venation dark reddish brown.

Legs. Integument black, partly dark red-brown. Vestiture dark brown, scopae blackish-brown (Fig. 6A).

Metasoma. Integument mostly black, red are the apical part of Tl, all of T2 except for a small black spot

anterior-medially, T3 except for a large black spot in the same position and apical margin of T4 (Fig.

6D). Tl and T2 on disc with long erect yellowish-white hairs; T3 - T4 covered with mostly blackish

and very short erect hairs; apical tergal hair bands on Tl - T4 medially broad, consisting of short sparse

white hairs; prepygidial and pygidial fimbriae black (Fig. 6A, D). Terga impunctate, finely shagreened

with a silky shine (Fig. 6D).

Male

Unknown.

Distribution

This species is only known from a single locality near Kamieskroon (Fig. 9).

Floral hosts

Unknown.

Seasonal activity

September.

Redivivoides karooensis sp. nov.

Figs 4, 7, 8

um:lsid:zoobank.org:act:00C81ElF-78C5-406D-8132-0CC6ABD24EB9

Diagnosis

Females of R. karooensis sp. nov. can be separated from other Redivivoides species by a combination

of the following characters: metasomal terga black to brown, T2 densely punctate with large punctures

and surface between punctures smooth and shiny, white apical tergal hair bands present (Fig. 7D),

prepygidial and pygidial fimbria dark brown to black (Fig. 7D). These characters also apply to males

(Fig. 8C) but the genitalia and S6 - S8 (Fig. 8D-H) should be checked to avoid potential confusion with

the unknown male of R. eardleyi.

Etymology

Named after the Karoo, the arid region in western and central South Africa where this species was found.

European Journal of Taxonomy 34 : 1-34 ( 2012 )

Type material (22 specimens)

Holotype

9 Sep. 1994, V.B. Whitehead / Phyllopodium heterophyllum / SAM-HYM-B009461“ (SAMC).

SOUTH AFRICA: W. Cape, Sauer, Suurfontein, 3218DC, 18°40’ E, 32°50’ S.

Paratypes

SOUTH AFRICA: 1 S, Farm Quaggaskop [3U19’ S, 18°39’ E], 29 Jul. 1992, M. Struck (SAMC);

1 $, 2 (?(?, Caries, Sarrisam Farm, 3017DA [30°40’ S, 17°40’ E], 11 Aug. 1994, V.B. Whitehead,

Phyllopodium pumilum (SAMC, RCMK); 1 $, Graafwater, 8 km West, 3218BA [32° 10’ S, 18°40’

E], 24 Sep. 1998, V.B. Whitehead, (SAMC); 3 ??, W. Cape, Rietvlei [32°09’ S, 18°46’ E], 282 m,

22 Sep. 2011, E. Packer (EPCT); 3 ??, 19.6 km S of Citrusdal [32°43’ S, 19°02’ E], 6 Sep. 1991, K.

Steiner, Phyllopodium heterophyllum (SAMC); 1 $, Sauer, Farm Suurfontein [32°50’ S, 18°33’ E], 13

Sep. 2001, V.B. Whitehead, Phyllopodium (SAMC); 4 Sauer, Suurfontein, 3218DC [32°50’ S,

18°40’ E], 25 Aug. 1994, V.B. Whitehead, Phyllopodium heterophyllum (SAMC); 1 $, 1 Sauer,

Suurfontein, 3218DC [32°50’ S, 18°40’ E], 1 Sep. 1994, V.B. Whitehead, Phyllopodium heterophyllum

(SAMC); 2 ??, 1 (?, Piketberg, Hartebeesrivier, 3218DC [32°50’ S, 18°40’ E], 23 Aug. 1991, V.B.

Whitehead, Polycarena (RCMK).

Fig. 7. Redivivoides karooensis sp. nov., $. A. Eateral view. B. Head. C. Scutum and scutellum.

D. Metasoma (dorsal view). Scale bar: 1 mm.

KUHLMANN M., Revision of the bee genus Redivivoides

Fig. 8. Redivivoides karooensis sp. nov., 3 . A. Lateral view. B. Seutum and seutellum. C. Metasoma

(dorsal view). D. S6 (ventral view). E. S7 (dorsal view). F. S8, apieal end (baekseattered eleetron

image). G. Genitalia (dorsal view). H. Genitalia (lateral view). Seale bar: 1 mm.

European Journal of Taxonomy 34 : 1-34 ( 2012 )

Description

Female

Body length. 9.5-10.5 mm

Head. Head slightly wider than long. Integument blaek exeept mandibles and labmm largely dark

reddish-brown. Faee sparsely eovered with long, yellowish-grey, ereet hairs intermixed with blaek hairs

espeeially along the inner eye margins and on vertex (Fig. 7B). Clypeus slightly eonvex, apieally narrowly

impunetate; medially densely eovered with medium-sized punetures that beeome gradually smaller

and denser towards the lateral and upper margins; surfaee between punetures shiny or superfieially

shagreened (Fig. 7B). Malar area medially narrow, almost linear. Antenna blaek, ventrally orange to

reddish-brown.

Mesosoma. Integument blaek. Mesoseutal dise between punetures smooth and shiny; dise densely

(i = 0.5-1.0 d) and finely punetate (Fig. 7C). Mesoseutum, seutellum, metanotum, mesepistemum and

propodeum eovered with long yellowish-brown ereet hairs, blaek hairs intermixed on mesoseutum

(Fig. 1C).

Wings. Yellowish-brown; wing venation brown.

Legs. Integument blaek to dark reddish-brown. Vestiture of femora whitish-yellow, on tibiae and tarsi

dark brown to blaek, seopae very sparse, dark brown to whitish-grey.

Metasoma. Integument blaek, apieal margins of terga narrowly translueent reddish-brown (Fig. 7D). T1

apieally and T2 on dise sparsely eovered with long ereet yellowish-white hairs; T3 - T4 eovered with

short yellow to blaek hairs; apieal tergal hair band missing on Tl, on T2 - T4 broadly white (Fig. 7A, D).

Prepygidial and pygidial fimbriae blaek. Tl almost impunetate, polished and shiny, T2 - T4 shiny, with

very fine, superfieial and sparse punetation that beeomes progressively denser on apieal terga (Fig. 7D).

Male

Body eength. 8.5-10.5 mm.

Head. Head slightly wider than long. Integument blaek exeept tip of mandible partly dark reddish-

brown. Faee densely eovered with long, yellowish-brown, ereet hairs intermixed with blaek hairs along

the inner eye margins and on vertex. Malar area medially narrow, almost linear. Antenna blaek, ventrally

yellowish to reddish-brown.

Mesosoma. Integument blaek. Mesoseutal dise between punetures smooth and shiny or superfieially

shagreened; dise densely (i = 0.5-1.0 d) and finely punetate (Fig. 8B). Mesoseutum, seutellum, meta¬

notum, mesepistemum and propodeum eovered with long yellowish-brown ereet hairs, on seutellum

blaek hairs intermixed.

Wings. Yellowish-brown; wing venation brown.

Legs. Integument blaek, tibiae and tarsi partly dark red-brown to yellowish-brown. Vestiture yellowish-

brown.

Metasoma. Integument blaek, apieal margins of terga narrowly translueent reddish-brown (Fig. 8C).

Tl - T3 eovered with long ereet yellowish-white hairs; T4 - T6 eovered with short blaek hairs; apieal

tergal hair band missing on Tl, on T2 - T5 narrow and sparse, white (Fig. 8A, C). Tl almost impunetate,

polished and shiny, T2 - T4 shiny, with very fine, superfieial and sparse punetation that beeomes

progressively denser on apieal terga (Fig. 8C).

KUHLMANN M., Revision of the bee genus Redivivoides

Terminalia. Genitalia (Fig. 8G-H), S6 (Fig. 8D), S7 (Fig. 8E) and terminal plate of S8 (Fig. 8F) as

illustrated.

Distribution

The species is known from a few places in southern Namaqualand and low lying areas west of the

Cedarberg Mountains (Fig. 4).

Floral hosts

Scrophulariaceae: Phyllopodium heterophyllum, Phyllopodium pumilum, Phyllopodium spec., Poly-

carena spec.

Seasonal activity

July - September.

Redivivoides namaquaensis sp. nov.

Figs 9, 10, 11

um:lsid:zoobank.org:act:AEF68F53-B181-4D12-8DlC-98QQ31A57F2E

Diagnosis

Males and females of R. namaquaensis sp. nov. can be separated from other Redivivoides species by

their black to brown metasomal terga in combination with dark brown to black prepygidial fimbria and

orange-brown pygidial fimbria (Figs lOD, IIC).

Etymology

Named after Namaqualand, the arid region in NW South Africa where this species occurs.

Type material (71 specimens)

Holotype

?, V.B. Whitehead, 16 Sep. 1999, Selago sp. /B009463” (SAMC). SOUTH AFRICA: N Cape, 31°46’

S, 20°03’ E, Middlepos, Elandsfontein.

Paratypes

SOUTH AFRICA: 5 $$, same date as holotype (SAMC); 4 $$, 2 SS, Nieuwoudtville Flower

Reserve, 3119AC [3U22’ S, 19°08’ E], 28 Aug. 1995, V.B. Whitehead, Selago sp. (SAMC, RCMK);

1 $, Nieuwoudtville Flower Reserve, 3119AC [31°22’ S, 19°08’ E], 5 Sep. 1995, V.B. Whitehead,

Selago sp. (SAMC); 1 (?, Nieuwoudtville Flower Reserve, 3119AC [3U22’ S, 19°08’ E], 30 Jul. 1997,

V.B. Whitehead, Selago sp. (SAMC); 1 S, Nieuwoudtville Flower Reserve, Glen Eyon [31°22’ S,

19°08’ E], 21 Jul. 1997, V.B. Whitehead, Selago pinguicola (SAMC); 2 SS, Nieuwoudtville Flower

Reserve, Glen Eyon [31°22’ S, 19°08’ E], 31 Jul. 1997, V.B. Whitehead, Selago pinguicola (SAMC);

3 SS, Nieuwoudtville Flower Reserve, Glen Eyon, 3119AC [31°22’ S, 19°08’ E], 14 Aug. 1997, V.B.

Whitehead, Selago pinguicola E. Mey. (SAMC); 1 S, Nieuwodtville Res., 3119AC [31°22’ S, 19°08’

E], 19 Aug. 1986, V.B. Whitehead, Othonna (SAMC); 1 S, N. Cape, Nieuwoudtville, Glen Eyon,

Renosterveld [31°24’42” S, 19°09’00” E], 700 m, 23 Aug. 2006, K. T imm erma rm (RCMK); 10

FarmBloemfontein, 3120CC [31°50’S, 20°10’E], 3 Oct. 1985, K. Steiner, (SAMC); XISS,

Farm Hartebeestfontein, 3120CC [31°50’ S, 20° 10’ E], 27 Aug. 1990, K. Steiner, Arctotheca calendula

(SAMC); 7 SS, Middlepos, Farm Hartebeestfontein, 3120CC [31°50’ S, 20°10’ E], 27 Aug. 1990, V.B.

Whitehead, Arctotheca, Polycarena (SAMC, RCMK); 1 S, 12 km N of Wuppertal, 3219AA [32°09’

S, 19°12’ E], 30 Aug. 1990, V.B. Whitehead, Polycarena (SAMC); 3 SS, 5 km S ofBidouw, 3219AA

[32°10’ S, 19°10’ E], 30 Aug. 1990, K. Steiner, Polycarena (SAMC); 9 Middelpos, Bloemfontein,

European Journal of Taxonomy 34 : 1-34 ( 2012 )

3219BB [32°10’ S, 19°50’ E], 3 Oct. 1985, V.B. Whitehead, Polycarena (SAMC, RCMK); 2 29,7

km NW Sutherland [32°13’ S, 20°30’ E], 27 Sep. 1985, V.B. Whitehead (SAMC); 1 ?, Wuppertal, 2

km N of, 3219AA [32°15’ S, 19°12’ E], 29 Aug. 1990, V.B. Whitehead, Polycarena (SAMC); 1 S,

Holfontein, 20 km S Clanwilliam [32°20’ S, 18°55’ E], 24 Aug. 1983, V.B. Whitehead, Hemimeris

racemosa (SAMC); 1 $, Farm Kanolfontein, 3220AD [32°20’ S, 20°20’ E], 22 Sep. 1985, K. Steiner,

Polycarena (SAMC); 1 S, Piketberg, Hartebeesrivier, 3218DC [32°50’ S, 18°40’ E], 23 Aug. 1991, V.B.

Whitehead, ^Polycarena (SAMC); 1 (J, Sauer, Suurfontein, 3218DC [32°50’ S, 18°40’E], 1 Sep. 1994,

V.B. Whitehead, Phyllopodium heterophyllum (SAMC).

Description

Female

Body length. 10.0-11.5 mm.

Head. Head slightly wider than long. Integument blaek exeept mandible and sometimes labrum partly

dark reddish-brown. Faee sparsely eovered with long, whitish-grey, ereet hairs intermixed with blaek

hairs along the inner eye margins and on vertex (Fig. lOB). Clypeus mostly flat, apieal margin almost

20E 25E 30E

Fig. 9. Distribution of Redivivoides eardleyi sp. nov., R. kamieskroonensis sp. nov. mdR. namaquaensis

sp. nov. in South Afriea.

KUHLMANN M., Revision of the bee genus Redivivoides

impunctate; medially densely covered with medium-sized punctures that become gradually smaller

and denser towards lateral and upper margins; surface between punctures shiny (Fig. lOB). Malar area

medially narrow, almost linear. Antenna black, ventrally dark orange to reddish-brown.

Mesosoma. Integument black. Mesoscutal disc between punctures smooth and shiny; disc densely

(i = 0.5-1.0 d) and finely punctate (Fig. IOC). Mesoscutum, scutellum, metanotum, mesepistemum and

propodeum covered with long yellowish-brown erect hairs, on the mesoscutum black hairs intermixed

(Fig. IOC).

Wings. Yellowish-brown; wing venation brown.

Legs. Integument black to dark reddish-brown. Vestiture of femora whitish-yellow, on tibiae and tarsi

dark brown to black, scopae yellowish-brown.

Metasoma. Integument dark brown to black, apical margins of terga narrowly translucent reddish to

yellowish-brown (Fig. lOD). T1 - T4 covered with erect yellowish-white hairs, becoming progressively

shorter from T1 to T4; apical tergal hair band sparse on Tl, on T2 - T4 broadly white (Fig. 10A,

D). Prepygidial fimbria dark brown to black, pygidial fimbria orange-brown. Terga impunctate, finely

shagreened and matt (Fig. lOD).

Fig. 10. Redivivoides namaquaensis sp. nov., $. A. Lateral view. B. Head. C. Scutum and scutellum.

D. Metasoma (dorsal view). Scale bar: 1 mm.

European Journal of Taxonomy 34 : 1-34 ( 2012 )

Fig. 11. Redivivoides namaquaensis sp. nov., S- A. Lateral view. B. Seutum and seutellum. C. Metasoma

(dorsal view). D. S6 (ventral view). E. S7 (dorsal view). F. S8, apieal end (baekseattered eleetron

image). G. Genitalia (dorsal view). H. Genitalia (lateral view). Seale bar: 1 mm.

KUHLMANN M., Revision of the bee genus Redivivoides

Male

Body length. 9.0-10.0 mm .

Head. Head slightly wider than long. Integument black except tip of mandible partly dark reddish-

brown. Face densely covered with long, whitish-grey, erect hairs intermixed with black hairs along the

inner eye margins and on vertex. Malar area medially narrow, a lm ost linear. Antenna black, ventrally

dark brown.

Mesosoma. Integument black. Mesoscutal disc between punctures smooth or superficially shagreened and

shiny; disc densely (i = 0.5 d) and very finely punctate (Fig. IIB). Mesoscutum, scutellum, metanotum,

mesepistemum and propodeum covered with long white to yellowish-grey erect hairs, partly intermixed

with a few black hairs.

Wings. Yellowish-brown; wing venation brown.

Legs. Integument black to reddish-brown. Vestiture whitish to yellowish-brown.

Metasoma. Integument dark brown to black, apical margins of terga narrowly translucent reddish to

yellowish-brown (Fig. IIC). T1 - T3 densely covered with long, erect yellowish-brown hairs, on T3

intermixed with black hairs; T4 - T6 covered with short black hairs; apical tergal hair band missing on

Tl, on T2 - T5 narrow and sparse, white; apical fringe of T7 orange (Fig. 11 A, C). Terga impunctate,

finely and densely shagreened, matt (Fig. IIC).

Terminalia. Genitalia (Fig. IIG-H), S6 (Fig. IID), S7 (Fig. HE) and terminal plate of S8 (Fig. IIF) as

illustrated.

Distribution

This species has the centre of its distribution on the Bokkeveld Plateau and in the Roggeveld Mountains

with a few records southwest of that area (Fig. 9).

Floral hosts

Asteraceae: Arctotheca calendula, Arctotheca spec., Othonna spec.; Scrophulariaceae: Hemimeris

racemosa, Phyllopodium heterophyllum, Polycarena spec., Selagopinguicola, Selago spec..

Seasonal activity

July - October.

Redivivoides simulans Michener, 1981

Figs 12, 13, 14

Redivivoides simulans Michener, 1981: 121-122, figs 80, 82-84 (p. 43), 97 (p. 48), Holotype $ (type

locality: Clanwilliam, South Africa) (Snow Entomological Museum, University of Kansas, Eawrence).

Diagnosis

Females of R. simulans can be separated from other Redivivoides species by a combination of the

following characters: metasomal terga at least partly red (at least on a narrow stripe on the apical part

of the discs of Tl - T3), metasomal terga with surface between punctures smooth and shiny (Fig. 12D),

scutum at least on the disc between punctures smooth and shiny (Fig. 12C), and head about as long as

wide (Fig. 12B). These characters also apply to males (Fig. 13C) but genitalia and S6 - S8 (Fig. 13D-H)

should be checked to avoid potential confusion with the unknown male of R. kamieskroonensis.

European Journal of Taxonomy 34 : 1-34 ( 2012 )

Both sexes of R. simulans were deseribed in detail by Miehener (1981).

Material examined

Speeimens examined (56 speeimens): SOUTH AFRICA: 2 $$,40 km N Port Nolloth [28°54’3” S,

16°44’3” E], 11 Sep. 2001 (CUIC, RCMK); 1 $, 1 ($, Skur&op St, 3118AB [31°10’ S, 18°20’ E],

18 Aug. 1983, V.B. Whitehead, white Hermannia prostrate (SAMC); 1 $, 1 ($, Farm Sarrisam,

3017DA [30°40’ S, 17°40’ E], 14 Aug. 1993, K. Steiner, Hermannia sp. (SAMC); 1 $, Ouberg Pass,

27 km SE Vamhynsdorp, Fynbos [31°48’07” S, 18°55’00” E], 380 m, 9 Aug. 2011, M. Kuhlmann

(RCMK); 2 $$, 2 S3, near Ratelfontein [31°56’ S, 18°3r E], 21 Sep. 2007, C. Eardley (SANC);

1 $, Koinonia [32°0r42” S, 18°39’34” E], 24 Sep. 2011, C. Eardley (SANC); 2 $ $, 1 ($, Ratelfontein

/ Olaf Bergfontein [32°02’ S, 18°35’ E], 7 Oet. 1995, F.W., S.K. & R.W. Gess, visiting yellow fls. semi-

prostrate Hermannia sp., Stereuliaeeae, 95/96/273 (AMGS); 2 $$, Trawal, 15 km SSW [32°02’ S,

18°35’ E], 21 Sep. 2007, T.E. Griswold (EMUS); 1 $, 1 ($, S Skur&op [32°03’26”S, 18°40’27” E], 24

Sep. 2011, C. Eardley (SANC); 1 $, Eamberts Bay [32°04’2F’ S, 18°22’12” E], 160 m, 13 Sep. 2005,

C. Eardley (SANC); 5 $$, 1 ($, 5 km N Clanwilliam [32°08’1” S, 18°5r3” E], 94 m, 20 Sep. 2001

(CUIC); 2 $$, 11 km N Clanwilliam [32°05’08” S, 18°49’25” E], 21 Sep. 2011, C. Eardley (SANC);

1 $, 20 mi. W. of Clanwilliam [32°09’ S, 18°36’ E], 19 Sep. 1966, C.D. Miehener (SAMC); 1 ($, 9 km

W Clanwilliam [32°10’1F’ S, 18°48’2F’ E], 22 Sep. 2011, C. Eardley (SANC); 1 $, 1 ($, Clanwilliam

Fig. 12. Redivivoides simulans Miehener, 1981, $. A. Eateral view. B. Head. C. Seutum and seutellum.

D. Metasoma (dorsal view). Seale bar: 1 mm.

KUHLMANN M., Revision of the bee genus Redivivoides

Fig. 13. Redivivoides simulans Michener, 1981, S- A. Lateral view. B. Seutum and seutellum.

C. Metasoma (dorsal view). D. S6 (ventral view). E. S7 (dorsal view). F. S8, apieal end (baekseattered

eleetron image). G. Genitalia (dorsal view). H. Genitalia (lateral view). Seale bar: 1 mm.

European Journal of Taxonomy 34 : 1-34 ( 2012 )

[32°10’ S, 18°53’ E], 8 Sep. 1966, M. & C.D. Michener (EMUS); 1 Droogerivier, Sandberg Sta.,

3218BC [32°18’ S, 18°37’ E], 24 Aug. 1987, V.B. Whitehead, Homeria (SAMC); 2 $$, 2 SS, Farm

Droogerivier, 3218BC [32° 18’ S, 18°37’ E], 8 Sep. 1987, K. Steiner, Lessertia rigida (SAMC); 1 3,

Droogerivier, Sandberg Sta., 3218BC [32°18’ S, 18°37’ E], 27 Aug. 1987, V.B. Whitehead, Homeria

(SAMC); 2 ??, 2 SS, Klein Alexandershoek to Het Kruis [32°20’ S, 18°46’ E], 26 Sep. 1985, F.W

& S.K. Gess (AMGS); 1 ?, Citrusdal [32°36’42” S, 18°59’49” E], 12 Sep. 2005, C. Eardley (SANC);

2 Sauer, 26 km W Piketberg [32°50’36” S, 18°33’58” E], 21 Get. 2008, C. Eardley (SANC); 5

$ Sauer, 26 km W Piketberg [32°50'37” S, 18°33’58” E], 21 Get. 2008, BNDanforth, on Hermannia

(CUIC); 4 ??, Sauer [32°50’ S, 18°34’ E], 21 Get. 2008, T.E. Griswold (EMUS); 1 ?, Atlantis,

3 km W, [33°34’ S, 18°27’ E], 150 m, 8-12 Get. 2002, M.E. Irwin, F.D. Parker (EMUS); 1 ?, Ganze-

kraal, 3318CB [33°40’ S, 18°20’ E], 22 Sep. 1999, K. Steiner, Nemesia affinis (SAMC).

Additional material

Additional reeords from the database of the South Afriean Museum, Cape Town (material in SAMC,

no data on number or sex of speeimens available): SGUTH AFRICA: Goegap Reserve, 4 km from

tarred road on eireular drive. Springbok [29°40’ S, 18°00’ E], 13 Aug. 1993, Whitehead, Zygophyllum;

20E 25E 30E

Fig. 14. Distribution of Redivivoides simulans Miehener, 1981 in South Afriea.

KUHLMANN M., Revision of the bee genus Redivivoides

Klipfontein [29°44’ S, 17°33’], 3 Aug. 1988, Whitehead, Zygophyllum, Bowesdorp, L. Namaqualand

[30°09’ S, 17°56’E], Sep. 1941, South African Museum; Grootvlei, Kamieskroon [30°10’ S, 17°50’E],

24 Aug. 1990, Whitehead, Indigofera; Farm Dik Nek on road to Hondeklipbaai [30° 19’ S, 17° 17’ E], 14

Aug. 1993, Whitehead; Farm Dik Nek [30°19’ S, 17°17’E] 130 m, 9 Aug. 1991, 14 Aug. 1993, Steiner

(SAMC); Diknek, Garies [30°19’ S, 17°17’ E], 18 Aug. 1995, Whitehead / Macpherson, Nemesia\

Wallekraal, Namaqualand [30°22’ S, 17°29’ E], Oct. 1950, South African Museum; Garies, Eittle

Namaqualand [30°33’ S, 17°58’ E], Jun. 1930, South African Museum; Garies, 9 km E. of Soutfontein

[30°33’ S, 17°58’ E], 22 Aug. 1990, Whitehead, Lebeckia; Kotzesrus, 12.6 km N. on rd to Groemivier

[30°58’ S, 17°49’ E], 12 Aug. 1997, Whitehead, Heliophila\ Kotzesrus, Hardevlei [30°58’ S, 17°49’ E],

28 Aug. 1997, Whitehead, Hermannia; Nieuwoudtville [31°22’ S, 19°07’ E], 31 Aug. 1985, Whitehead

/ Macpherson, Lotonotis serpens', Garies, Farm Sariesaam [30°40’ S, 17°40’ E], 14 Aug. 1993,

Steiner; 6.5 km N of Klawer t/o on N7 [31°45’ S, 18°39’ E], 6 Aug. 1989, Steiner, Homeria minuata',

Skurfkop station [31°56’ S, 18°36’ E], 18 Aug. 1983, Whitehead, Hermannia', Eamberts Bay [32°05’ S,

18°18’E],28Sep. 1977, Whitehead; Farm Skarpdam[32°05’S, 18.22’E],9Sep. 1987, Steiner/Whitehead,

Nemesia', 12 kmN of Clanwilliam [32°05’ S, 18°52’ E], 11 Aug. 1985, Whitehead, Hermannia', 11 kmN

of Clanwilliam [32°06’ S, 18°52’ E], 21 Aug. 1985, Whitehead / Macpherson; Kompagniesdrif, 19 km

E. of Eamberts Bay [32°08’ S, 18°29’ E], 14 Sep. 1984, Whitehead, Hermannia', Graafwater [32°08’ S,

18°35’ E], Oct. 1947, South African Museum Expedition; 10 km E of Bidouw Valley [32°08’ S, 19°19’

E], 8 Sep. 1985, Picker; Clanwilliam [32°10’ S, 18°53’ E], 11 Aug. 1984, Picker; Ramskop campsite,

Clanwilliam [32° 10’ S, 18°53’ E], 22 Aug. 1984, Whitehead / Macpherson, Trachyandra muricata',

Eeipoldtville, Elands Bay [32°12’ S, 18°28’ E], Oct. 1947, South African Museum Expedition; Elands

Bay [32°17’ S, 18°20’ E], 20 Sep. 1988, Whitehead, Polycarena seluginoides'. Elands Bay [32°17’ S,

18°20’ E], 26 Sep. 1985, Whitehead, Hermannia', Farm Skerpheuwel [32°18’ S, 18°20’ E], 28 Aug.

1987, Steiner, Nemesia', Farm Droogerivier [32°18’ S, 18°37’ E], 8 Sep. 1987, Steiner; Droogerivier

(Droerivier), Sandberg sta. [32° 18’ S, 18°37’ E], 27 Aug. 1987, Whitehead, Homeria', 11 Aug. 1988,

Whitehead, Cysticapnos', 8 Sep. 1987, Steiner, Lessertia rigida'. Farm Droogerivier (Droerivier)

[32°18’ S, 18°37’ E], 27 Aug. 1987, 8 Sep. 1987, Steiner, Lessertia rigida / Cysticapnos vesicarius',

Paleisheuwel, Alexanderhoek [32°18’ S, 18°43’ E], 12 Sep. 1991, Whitehead, Cysticapnos', Holfontein,

20 km S of Clanwilliam [32°20’ S, 18°55’ E], 29 Aug. 1984, Whitehead / Steiner, Hermannia', 28.8

km N of Citrusdal t/o [32°20’ S, 19°00’ E], 8 Aug. 1984, Whitehead, Hermannia', Farm Klawervlei,

23 km south of Clanwilliam [32°23’ S, 18°56’ E], 20 Aug. 1984, Whitehead, Hermannia, Lachenalia

mutabilis', Koraanshoogte, 13.5 km north of Citrusdal t/o [32°28’ S, 19°00’ E], 20 Aug. 1984, Whitehead

/ Macpherson, Hermannia', Oorwinningsfontein, 33 km N of Piketberg [32°36’ S, 18°44’ E], 30

Aug. 1987, Whitehead, Hermannia', 30 Jul. 1987, Whitehead, Hermannia', Eendekuil, Aroenfontein

[32°40’ S, 18°53’ E], 19 Aug. 1992, Whitehead; 1 km S of Droeryskloof Sta., Eendekuil [32°40’ S,

18°53’ E], 30 Aug. 1986, Whitehead, Hermannia, Sauer, Suurfontem [32°50’ S, 18°40’ E], 1 Sep. 1994,

Whitehead / Macpherson, Nemesia', Piketberg [32°54’ S, 18°44’ E], 21 Sep. 1985, Whitehead; Berg River,

Jantjiesfontem [33°05’ S, 18°25’ E], 25 Sep. 1991, Whitehead; Hopefield, Jantjiesfontem [33°05’ S,

18°25’ E], 17 Sep. 1991, Whitehead, Indigofera', 12 km SE of Yzerfontein t/o [33°20’ S, 18°17’ E], 28

Sep. 1988, Whitehead, Nemesia versicolor, between Darling and Yzerfontein [33°20’ S, 18°17’ E], 18

Sep. 1986, Steiner, Heliophila', Montagu Rocks, Bainskloof [33°35’ S, 19°08’E], 31 Sep. 1990, Steiner,

Moraea ramosissima, Ganzekraal [33°40’ S, 18°20’ E], 22 Sep. 1999, Steiner, Nemesia affinis'. Darling

road, 18 Sep. 1986, Whitehead, Heliophila.

Distribution

This is the most common and widespread of all Redivivoides species and has been found in most parts

of the winter rainfall area (Fig. 14).

European Journal of Taxonomy 34: 1-34 (2012)

Floral hosts

Asphodelaceae: Trachyandra muricata; Brassicaceae: Heliophila spec.; Fabaceae: Indigofera spec.,

Lebeckia spec., Lessertia rigida, Lotonotis hirsuta (= serpens)', Fumariaceae: Cysticapnos vesicaria',

Hyacinthaceae: Lachenalia mutabilis', Iridaceae: Moraea miniata, M. ramosissima, Moraea spec.;

Scrophulariaceae: Nemesia affinis, N. versicolor, Nemesia spec., Polycarena phyllopodioides

(= seluginoides)', Sterculiaceae: Hermannia spec.; Zygophyllaceae: Zygophyllum spec.

Seasonal activity

(June) July - October.

Redivivoides variabilis sp. nov.

Figs 15, 16, 17

um:lsid:zoobank.org:act:8353A5DA-3526-4FDC-A7EF-E31Q70DB45EF

Diagnosis

Females and males of R. variabilis can be separated from other Redivivoides species by their black to

brown metasomal terga which are smooth and shiny between punctures and that lack white apical hair

bands (Figs 15E-F, 16E). This is the only Redivivoides species where terga have an intense oily bluish

shine.

Etymology

The species is named after the colour variation of the females.

Type material (31 specimens)

Holotype

$, 17 Aug. 1999, K. Steiner / Host Plant: Melolobium obcordatum / SAM-HYM-B009467 (SAMC).

SOUTH AFRICA: N.C. Prov., Grootvlei, 3017BB, 17°50’E, 30°10’S.

Paratypes

SOUTH AFRICA: 1 ?, 1 (? same date as holotype (SAMC); 1 ?, 1 (?, Springbok [29°40’ S, 17°52’

E], 7 Sep. 1966, C.D. Michener (SAMC); 3 $$, Namaqualand, Springbok, Goegap Nature Res. (hills)

[29°4U S, 17°59’ E], 8-10 Sep. 1992, F.W. & S.K. Gess, on yellow fls. of Lebeckia spinescens Harv.

(Papilionaceae), 92/93/41, (AMGS); 2 $ Dassiefontein Farm, 14 km E Kammieskroon (sic) [30°09’

S, 17°59’ E], 16-17 Sep. 2001, C. Eardley (SANC); 1 ?, Skilpad, 3017BB [30°10’ S, 17°50’ E], 19

Aug. 1999, K. Steiner, Nemesia anisocarpa (SAMC); 1 $, Kamieskroon, Dassiefontein, 3017BB

[30°10’ S, 17°50’ E], 7 Sep. 1986, K. Steiner, Lotononis (SAMC); 1 ?, Farm Grootvlei, 3017BB

[30°10’ S, 17°50’ E], 24 Aug. 1990, K. Steiner, Eegume (SAMC); 1 S, Grootvlei, 20 km S Kamieskroon

[30° 10’ S, 17°50’ E], 10 Aug. 1984, V.B. Whitehead, yellow legume (SAMC); 2 $ ^ km E Karkams,

3017BD [30°20’ S, 17°50’ E], 10 Aug. 1985, V.B. Whitehead, Nemesia & Lachenalia (SAMC); 1 ?,

6.2 km NE of Karkams, 3017BD [30°20’ S, 17°50’ E], 4 Sep. 1986, K. Steiner, Nemesia (SAMC); 1 ?,

Garies, Welkom Farm, 3018CA [30°40’ S, 18°10’ E], 30 Sep. 1988, V.B. Whitehead, Lotononis serpens

(SAMC); 1 $, N. Cape, Nieuwoudtville, Flower Reserve East [31°21’56” S, 19°08’52” E], 735 m, 10

Aug. 2004, M. Kuhlmann (RCMK); 1 $ (black form), N. Cape, Nieuwoudtville, Flower Reserve East

[31°21’56”S, 19°08’52”E],735m,6Sep. 2002, M. Kuhlmann (RCMK); 1 ? (black form), 1 Cape,

Nieuwoudtville, Flower Reserve East [31°21’56” S, 19°08’52”E], 735 m, 3 Sep. 2003, M. Kuhlmann, on

Oxaliss^QC. (RCMK); 1 5,Nieuwoudtville, FarmGlenlyon, 3117AC [31°21’S, 19°08’E], 14Aug. 1997,

V.B. Whitehead, Lotononis serpens (SAMC); 1 S, Nieuwoudtville Flower Reserve [31°22’ S, 19°08’ E],

2 Aug. 1984, V.B. Whitehead, Lotononis serpens (SAMC); 2 5 ?, W. Cape, 24 km N of Garies [30°23’

S, 17°54’ E], 777 m, 25 Sep. 2011, E. Packer (EPCT, RCMK); 1 S, N. Cape, Nieuwoudtville, Glen

KUHLMANN M., Revision of the bee genus Redivivoides

Fig. 15. Redivivoides variabilis sp. nov., A. Lateral view (light form). B. Lateral view (black form).

C. Head. D. Scutum and scutellum. E. Metasoma (dorsal view) (light form). F. Metasoma (dorsal view)

(black form). Scale bar: 1 mm.

European Journal of Taxonomy 34: 1-34 (2012)

Fig. 16. Redivivoides variabilis sp. nov., S- A. Lateral view. B. Seutum and seutellum. C. Metasoma

(dorsal view). D. S6 (ventral view). E. S7 (dorsal view). F. S8, apieal end (baekseattered eleetron

image). G. Genitalia (dorsal view). H. Genitalia (lateral view). Seale bar: 1 mm.

KUHLMANN M., Revision of the bee genus Redivivoides

Lyon, Renosterveld [31°24’03” S, 19°08’34” E], 700 m, 18 Aug. 2004, M. Kuhlmann (RCMK); 1 $

(black form), Keiski Mts., 5 km S Farm Nooiensrivier, dolerite hill [31°45’54”S, 19°50’17” E], 1270

m, 29 Aug. 2010, M. Kuhlmann (RCMK); 1 5 (black form), Keiski Mts., 5 km S Farm Nooiensrivier,

dolerite hill [31°45’54” S, 19°50’17” E], 1270 m, 4 Sep. 2010, M. Kuhlmann (RCMK); 1 $, Skaapdam,

11 km E Eamberts Bay [32°05’ S, 18°22’E], 9 Sep. 1987, V.B. Whitehead, Nemesia (SAMC); 1 $ (black

form), Clanwilliam, Ramskop [32°10’ S, 18°53’ E], 21 Aug. 1984, V.B. Whitehead & M. Macpherson,

Homeria minuta (SAMC).

Description

Female

Body LENGTH. 12.0-13.0 mm.

Head. Head slightly wider than long. Integument black except median part of mandible and sometimes

labrum partly dark reddish-brown. Face sparsely covered with long, whitish-grey (in the dark form

all black. Fig. 15B), erect hairs intermixed with black hairs along the inner eye margins and on vertex

20E 25E 30E

Fig. 17. Distribution of Redivivoides variabilis sp. nov. in South Africa.

European Journal of Taxonomy 34: 1-34 (2012)

(Fig. 15C). Clypeus mostly flat, apically almost impunctate, punctures here large and seattered beeoming

rapidly smaller and denser towards the upper edge; surfaee between punetures shiny (Fig. 15C). Malar

area medially narrow, almost linear. Antenna blaek, ventrally sometimes partly dark reddish-brown.

Mesosoma. Integument blaek, tegulae partly yellowish-brown. Mesoseutal dise between punetures

smooth and shiny; dise densely (i = 0.5-1.0 d) and flnely punetate (Fig. 15D). Mesoseutum, seutellum,

metanotum, mesepistemum and propodeum eovered with long yellowish-brown ereet hairs, on the dise

of mesoseutum and seutellum some blaek hairs intermixed (in the dark form all blaek. Fig. 15B).

Wings. Yellowish-brown; wing venation brown.

Legs. Integument blaek to yellowish-brown. Vestiture yellowish-brown, seopae yellowish-brown. In

the dark form hairs of femora and most of tibiae blaek or dark brown, on tarsi yellowish-brown, seopae

darker yellowish-brown (Fig. 15B).

Metasoma. Integument blaek, exeept apieal tergal margins partly narrowly brownish translueent; terga

with intense oily bluish shine (Fig. 15E-F). T1 apieally sparsely eovered with a few long ereet yellowish

hairs; T2 - T4 eovered with short yellowish-white hairs; apieal tergal hair band missing on Tl, on

T2 - T4 very sparse and only laterally developed (Fig. 15A-E). In the dark form all hairs blaek, no

apieal tergal hair bands (Fig. 15B-F). Prepygidial fimbria dorsally blaek, laterally partly yellowish white

(in dark form all blaek), pygidial fimbria blaek. Terga densely but flnely punetate, smooth and shiny

between punetures (Fig. 15E-F).

Male

Body EENGTH. 12.0-13.0 mm.

Head. Head slightly wider than long. Integument blaek exeept tip of mandible partly dark reddish-brown.

Faee densely eovered with long, whitish-grey to yellowish-brown, ereet hairs intermixed with blaek

hairs along the inner eye margins and on vertex. Malar area medially narrow, a lm ost linear. Antenna

blaek, ventrally dark reddish-brown.

Mesosoma. Integument blaek, tegulae partly yellowish-brown. Mesoseutal dise between punetures

smooth and shiny; dise densely (i = 0.5-1.0 d) and flnely punetate (Fig. 16B). Mesoseutum, seutellum,

metanotum, mesepistemum and propodeum eovered with long yellowish-brown ereet hairs, on the dise

of mesoseutum and seutellum some blaek hairs intermixed (Fig. 16B).

Wings. Yellowish-brown; wing venation brown.

Eegs. Integument blaek, tibiae and tarsi partly yellowish-brown. Vestiture yellowish-white.

Metasoma. Integument blaek, exeept apieal tergal margins partly narrowly brownish translueent; Tl -

T3 with intense oily bluish shine, T4 - T5 less so (Fig. 16A, C). Tl eompletely and T2 on dise eovered

with long ereet yellowish-white hairs; T2 - T3 densely eovered with short ereet yellowish-white either

yellowish-white or blaek on T4 and blaek on T5 - T6; apieal tergal hair band missing on Tl, on T2 - T5

sparse and narrow, yellowish-white to white (Fig. 16A, C). Terga densely but flnely punetate, smooth

and shiny between punetures (Fig. 16C). T7 with pygidial plate that is redueed to a narrow longitudinal,

slightly elevated and shiny ridge.

Terminaeia. Genitalia (Fig. 16G-H), S6 (Fig. 16D), S7 (Fig. 16E) and terminal plate of S8 (Fig. 16F) as

illustrated.

KUHLMANN M., Revision of the bee genus Redivivoides

Distribution

The species has been found in most parts of Namaqualand to the northern parts of the Roggeveld

Mountains in the southeast (Fig. 17). The colour form with females all black is only known from the

higher grounds of the south-eastern part of the species' range while the females with light brown pilosity

mostly occur in lower areas and the north (Fig. 17). Colour variation is also known from Rediviva gigas

Whitehead & Steiner, 1993 and R. parva Whitehead & Steiner, 2001. In R. gigas there is no obvious

geographical pattern of colour variation but the black form of R. parva seems to be concentrated in the

north and in lower lying areas in the west of its range (Whitehead & Steiner 2001).

Floral hosts

Fabaceae: Lebeckia spinescens, Lotonotis hirsuta (= serpens), Lotononis spec., Melolobium obcor-

datum', Hyacinthaceae: Lachenalia spec.; Iridaceae: Moraea miniata, Oxalidaceae: Oxalis spec.;

Scrophulariaceae: Nemesia anisocarpa, Nemesia spec.

Seasonal activity

August - September.

Key to Redivivoides species

Females

1. Metasomal terga extensively and brightly red (Figs 2D, 6D, 12D), rarely only on a narrow stripe

of the apical part of the discs of T1 - T3 . 2

-Metasomal terga black or brown (Figs 5D, 7D, lOD, 15E-F), sometimes apical tergal margins

narrowly translucent dark reddish brown.4

2. Scutum impunctate, shagreened and matt (Fig. 2C). R. capensis sp. nov.

- Scutum finely punctate and at least on disc between punctures smooth and shiny (Figs 6C, 12C) ... 3

3. Metasomal terga finely shagreened and with a dull shine (Fig. 6D); head distinctly wider than long

(Fig. 6B). R. kamieskroonensis sp. nov.

-Metasomal terga between punctures smooth and shiny (Fig. 12D); head only slightly wider than

long (Fig. 12B). R. simulans MichQnQX, 1981

4. Metasomal terga finely shagreened and dull (Fig. lOD); prepygidial and pygidial fimbria yellowish

brown (Fig. lOD) . R. namaquaensis sp. nov.

-Metasomal terga between punctures smooth and shiny (Figs 5D, 7D, 15E-F); prepygidial and

pygidial fimbria dark brown to black (Figs 5D, 7D, 15E-F) . 5

5. Metasomal terga without white apical tergal hair bands (Fig. 15E-F); terga with intense oily

bluish shine . R. variabilis sp. nov.

- Metasomal terga with white apical tergal hair bands (Figs 5D, 7D); terga without oily bluish shine ... 6

6. Disc of T2 sparsely punctate (i = 2-3 d); punctures minute (Fig. 5D); head and clypeus as in Fig.

5B . R. eardleyi sp. nov.

-Disc of T1 densely punctate (i = d); punctures larger (Fig. 7D); head and clypeus as in Fig. 7B

. R. karooensis sp. nov.

European Journal of Taxonomy 34: 1-34 (2012)

Males

The males of R. eardleyi sp. nov. and R. kamieskroonensis sp. nov. are unknown.

1. Metasomal terga extensively and brightly red, rarely only on a narrow stripe of the apieal part of the

dises of T1 - T3 (Figs 3C, 13C). 2

- Metasomal terga blaek or brown, sometimes apieal tergal margins narrowly translueent dark reddish

brown (Figs 8C, IIC, 16C). 3

2. Seutum impunetate, shagreened and matt (Fig. 3B); S6 - S8 and genitalia as in Fig. 3D-H

. R. capensis sp. nov.

- Seutum finely punetate and at least on dise between punetures smooth and shiny (Fig. 13B); S6 -

S8 and genitalia as in Fig. 13D-H . R. simulans Miehener, 1981

3. Metasomal terga finely shagreened and dull (Fig. IIC); prepygidial and pygidial fimbria yellowish

brown (Fig. IIC); S6 - S8 and genitalia as in Fig. IID-H. R. namaquaensis sp. nov.

-Metasomal terga between punetures smooth and shiny at least on T1 - T2 (Figs 8C, 16C); prepy¬

gidial and pygidial fimbria brown to blaek (Figs 8C, 16C); S6 - S8 and genitalia different.4

4. Metasomal terga with oily bluish shine; S6 - S8 and genitalia as in Fig. 16D-H.

. R. variabilis sp. nov.

- Metasomal terga without oily bluish shine; S6 - S8 and genitalia as in Fig. 8D-H.

. R. karooensis sp. nov.

Discussion

Redefinition of the genus Redivivoides

Within the Melittinae Miehez et al. (2009), based on a phylogenetie study, reeognize two tribes; the

Maeropidini eomprises genera with two submarginal eells (Afrodasypoda, Macropis, Promelitta and the

fossil taxa Eomacropis and Paleomacropis) and the Melittini, eomprising genera with three submarginal

eells {Melitta, Rediviva, Redivivoides). In the Melittini Melitta is eharaeterized by a well-defined, large

and dull propodeal triangle (exeept Melitta schultzei; Miehez & Eardley 2007) while in Rediviva and

Redivivoides this is ill-defined, small and shiny, although in some speeies superfieially seulptured.

In both Macropis (Fig. lA-B) and Rediviva (Fig. IC-D) the females ean easily be reeognized by their

broadened hind tibia and basitarsus with the seopa eonsisting of densely plumose, velvety pubeseenee

for transporting fioral oil, while the elongate fore tarsi (extreme in some speeies) are a synapomorphy

of Rediviva (Miehener 1981, 2007). In eontrast the females of Redivivoides have short front legs with

simple hairs and a slender hind tibia, as in Melitta (Miehener 1981). The seopa eonsists of sparse simple

bristles (more dense and usually with at least a few shortly branehed hairs in Melitta, Fig. IG) with

the anterior margin of the seopa shining, unlike in other genera (Fig. IE) (Miehener 1981). The hind

basitasus of Redivivoides females is slightly broadened, about 2-2.5 times as long as its maximum width

with the posterior margin eurved and without an apieal proeess (Fig. IF) while in Melitta it is slender,

three or more times as long as broad, parallel-sided and with a small apieal proeess (Fig. IH) (Miehener

1981). In Redivivoides a penieillus is generally missing while it is present in all Melitta speeies known

to me and modified into a brush-like strueture used for lining the wall of brood eells in Rediviva and

Macropis (Sehaffier & Dotterl 2011).

As in females, male Melitta ean be reeognized by the large and dull propodeal triangle (exeept Melitta

schultzei) (ill-defined, small and shiny in Rediviva and Redivivoides), S7 having a large plate that is

truneate or emarginate apieally (sometimes with hair patehes at posterior lateral angles) and with lobes

KUHLMANN M., Revision of the bee genus Redivivoides

or processes reduced to small sclerotic structures like in Melitta budensis (Michez & Eardley 2007). In

Rediviva and Redivivoides SI in turn has a small disc with the apex bifid and/or with membranous lobes

and the gonostyli are relatively long (Michener 1981, 2007).

The differentiation of Rediviva ?ind Redivivoides males is problematic. Michener (1981,2007) suggested

that the broad, subtrunctate apex of the male SI, with its large laterally attached, vertical, membranous

hairy lobes (Fig. 13E) is a unique apomorphy of Redivivoides. However, these characters seem to be

species specific for R. simulans and they are either absent {R. karooensis, R. namaquaensis) (Figs 8E,

HE) or reduced {R. capensis, R. variabilis) (Figs 3E, 16E) in the other four species (the males of

R. eardleyi and R. kamieskroonensis are unknown). Michener (1981) mentioned the weak pygidial

plate in male R. simulans as a character differentiating Redivivoides from both Melitta and Rediviva.

A pygidial plate is also present in male Macropis and in R. variabilis, where it is reduced to a narrow

longitudinal, slightly elevated and shiny ridge but it is missing in all the other three species making it

unsuitable as a differentiating character.

Generally, the morphology of S7, S8 and the genitalia of Redivivoides males (Figs 3, 8, 11, 13, 16) refiect

much of the morphological diversity known from the much more species-rich Rediviva as illustrated in

Whitehead & Steiner (2001) and Whitehead et al. (2008). Although well characterized as species, the

obvious lack of apomorphic characters defining the males of Redivivoides at the genus level make

their identification difficult and indicate the close relationship of both genera as suggested by Michener

(1981) and Michez et al. (2009).

Origin of oil-collecting

The origin of oil-collecting in bees has been much debated and this behaviour has evolved independently

at least five times in the Centridini, Ctenoplectrini, Exomalopsini, Tetrapediini andMelittinae (Buchmann

1987; Vogel 1974, 1986, 1990; Renner & Schaefer 2010). Within the Melittinae fioral oil is collected by

bees of the gonom Macropis md Rediviva (Michener 2007). Melittids are the most basal bees (Danforth

et al. 2006) and, thus, they can be a key-group for understanding the evolution of oil-collecting in bees

in general. Michener (1981) and later Michez et al. (2009), based on the results of a phylogenetic study,

discussed whether oil-collecting has evolved only once in melittids or if there are two independent

origins in Macropis and Rediviva.

In both scenarios the phylogenetic position of Redivivoides is crucial to understanding the evolutionary

history of early bees. In Michez et a/.’s (2009) analysis Redivivoides arose from Rediviva, making the

latter paraphyletic, indicating a loss of oil-collecting in Redivivoides. However, only two of the currently

24 described Rediviva species and only R. simulans of the now seven known Redivivoides species were

included in the study. A broader approach including more species is required to confirm the paraphyly

of Rediviva.

If Redivivoides evolved from Rediviva the question is why Redivivoides is only present in the winter

rainfall region in the west of South Africa whereas Rediviva also occurs in the summer rainfall area in

the east of the country. Either Redivivoides evolved in the winter rainfall region and has not spread to

the east of South Africa or it became extinct there. An interesting parallel case is the 11 of 35 described

species of the Colletes fasciatus-growp in western South Africa that have a reduced scopa, which is

unique in the genus (Kuhlmann 2006, 2007).

Based on the present taxonomic revision there is now additional evidence that Redivivoides species might

have lost their oil-collecting structures, as presumably happened in some Neotropical Centris species

(Michener 2007). With the exception of Redivivoides all non oil-collecting genera of the Melittinae

have a slender hind basitarsus and a dense scopa with at least some branched long hairs. In contrast, the

European Journal of Taxonomy 34: 1-34 (2012)

hind tibia and basitarsus of oil-collecting Macropis and Rediviva is very broad and the scopa consists of

densely plumose, short pilosity for oil-collecting embedded in a matrix of sparse, long, simple, emergent

hairs (Fig. lA-D) (Michener 1981), with the latter reminiscent of Redivivoides (Fig. lE-F). Thus, the

broadened and posteriorly curved hind basitarsus lacking an apical process (present in Melitta, Macropis

and in modified form in some Rediviva), the sparse scopa consisting of only simple hairs in Redivivoides

and the lack of a penicillus (present in Melitta and modified into a brush-like structure in Rediviva and

Macropis) might be remnants indicating a loss of oil-collecting structures. This view is supported by the

phylogenetic position of R. simulans, arising from Rediviva, rendering Rediviva paraphyletic (Michez

et al. 2009), the similar structure of male genitalia, S7 and S8 and a study of fore and mid leg anatomy

by Steiner & Cruz (2006) suggesting that Redivivoides has descended from an oil-collecting ancestor.

The close relationship of Redivivoides and Rediviva is also refiected in fiower visitation. Most Rediviva

collect fioral oil on Scrophulariaceae (Whitehead & Steiner 2001; Whitehead et al. 2008) and pollen

from this plant family is also a major constituent of female scopal pollen loads (Kuhlmann unpubl.)

but it is rarely used by other Melittidae (Michez et al. 2008). Whilst no information is available on

fiower visitation of R. eardleyi sp. nov. and R. kamieskroonensis sp. nov. and only a single observation

for R. capensis sp. nov. all the other Redivivoides species were found visiting Scrophulariaceae with

R. karooensis sp. nov. and R. namaquaensis sp. nov. obviously having a preference for this family.

Although Kuhlmann & Eardley (2012) showed that Scrophulariaceae are visited and their pollen

collected by a range of bee species it is usually only a minor component in scopal pollen loads (Kuhlmann

unpubl.). Thus, the shared preference for Scrophulariaceae even though only based on a small number of

observations can be seen as another indicator for the common ancestry of both genera.

However, a robust phytogeny including more species of Rediviva and Redivivoides is required to better

understand the evolution of oil-collecting and phylogeography of both genera.

Acknowledgements

1 am very much indebted to the following colleagues (in alphabetical order of collection acronyms)

who made specimens under their curation available for this study: S.K. Gess and F.W. Gess (AMGS),

B.N. Danforth (CUIC), T. Griswold (EMUS), E. Packer (EPCT), D. Earsen and S. van Noort (SAMC)

and C.D. Eardley (SANG). Northern Cape Nature Conservation Service and the Western Cape Nature

Conservation Board (CapeNature) are gratefully acknowledged for giving their permission to collect

bees. Gavin Broad, Eondon, kindly made linguistic corrections. The insightful comments of two

anonymous reviewers helped to improve the paper.

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Manuscript received: 30 July 2012

Manuscript accepted: 7 December 2012

Published on: 19 December 2012

Topic editor: Koen Martens

Printed versions of all papers are also deposited in the libraries of the institutes that are members of

the EJT eonsortium: Museum National d’Histoire Naturelle, Paris, Franee; National Botanie Garden

of Belgium, Meise, Belgium; Royal Museum for Central Alfiea, Tervuren, Belgium; Natural History

Museum, Eondon, United Kingdom; Royal Belgian Institute of Natural Seienees, Brussels, Belgium;

Natural History Museum of Denmark, Copenhagen, Denmark.