Phylogenetic analyses of tribe Anthidiini

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UNIVERSITY OF CAPE COAST

SCHOOL OF BIOLOGICAL SCIENCES

PHYLOGENETIC ANALYSES OF THE BEE TRIBE ANTHIDIINI AND REVISION OF THE AFROTROPICAL GENUS

Anthidiellum (COCKERELL)

BY

ROFELA COMBEY

Thesis submitted to the Department of Entomology and Wildlife of the School of Biological Sciences, University of Cape Coast in partial

fulfillment of the requirements for the award of Doctor of Philosophy in Zoology

JUNE 2008

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DECLARATION

Candidate’s declaration

I hereby declare that this thesis is the result of my own original work

and that no part has been presented for another degree in this

university or elsewhere.

Candidate’s Signature:_____________________

Name: Rofela Combey …………………….Date:_________________

Supervisors’ Declaration

We hereby declare that the preparation and presentation of the thesis

were supervised in accordance with the guidelines on supervision of

thesis laid down by the University of Cape Coast.

Principal Supervisor’s Signature:____________________

Name: Dr. Peter Kofi Kwapong Date:_____________

Co-Supervisor’s Signature:__________________________

Name: Professor (Mrs) Mary Botchey Date:____________

Co-Supervisor’s Signature:___________________

Name: Dr. Connal Desmond Eardley Date:___________

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CHAPTER ONE

INTRODUCTION

The Concept of Phylogeny

There are so many different kinds of organisms on Earth that are

believed to exist as a result of evolution. If this evolutionary history or the

phylogeny of an organism is traced back, it connects through shared ancestors

to lineages of other organisms (Lipscomb, 1998). The scientific method that

analysizes and groups organisms that share derived characters is called

cladistic or phylogenetic systematics. This important aspect of systematics

was first published by Henning (1966) who argued that only shared derived

characters could possibly give us information about phylogeny.

The principle behind phylogenetics is that, organisms should be

classified according to their evolutionary relationships, and these relationships

are analysized based on characters occurring in either the primitive or advance

state. Primitive (plesiomorphic) characters are those characters in which all

members of the taxa possess. However these characters do not reveal the

relationship within a taxa. Advanced (apomorphic) characters on the other

hand are derived traits which appear in some members of the taxa and reveal

phylogenetic relationships among the taxa (groups). Taxa that share many

derived characters are grouped more closely together than those that do not

share similarly derived characters. The relationships are shown in a branching

hierarchical tree called a cladogram (Lipscomb, 1998; Michener, 2000).

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This phylogenetic tree or cladogram provides the basis of

evolutionary interpretation; it also provides the classification by which other

biologists communicate information about an organism; the cladogram and its

corresponding classification predicts properties of newly discovered or poorly

known organisms (Lipscomb, 1998; Michener, 2000).

Why Bees?

Among the hymenopteran insects, bees occupy a unique position in the

world of Arthropods. They are entwined into most aspects of human culture

and mythology, not to mention our agriculture, economy and general ecology.

Several centuries ago, bees were used to instil in man ideal notions of

industriousness, loyalty, obedience and sacrifice not through denigration but

through almost holy reverence to a greater good. Even the famous parable

from the Bible teaching us to “go to the ant” originally included reference to

the labour of bees (“Go to the bee, and learn how diligent she is, and what a

noble work she produces, whose labours kings and private men use for their

use, she is desired and honoured by all, and though weak in strength she

values wisdom and prevails:” Proverbs, Chapter 6, Septuagint version-i.e.,

oldest Greek version and perhaps the oldest extant version of the Holy Bible).

It is believed that such emotions and concepts connected to bees flow through

peoples’ common natural heritage; even the great “Mother Goddess”

uncovered at the root of most Western religions were symbolized by bees

(Engel, 2000, 2001).

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Today, bees are known by far to be major contributors to essential

ecological services including pollination. Globally, pollination and pollinator

issues have occupied keystone position in maintenance of biodiversity, in both

natural and agro-ecosystem. Pollination services provided by most bee

pollinators appear to be different from most other essential ecosystem services

in that, it is biodiversity dependent; for example many plant species require

specific pollinators and similarly many pollinator species require specific food

plant (Constanza, 1987; O’Toole, 1993; Kevan, et. al., 2002 and Fact sheet,

2004). Bees are almost widely used in pollination management in agriculture

(Prescott-Allen and Prescott-Allen, 1990; Ingram et. al., 1996; Kenmore and

Krell, 1998). Aside this essential service, hive products provide enormous

economic value to mankind.

In view of these essential roles played by bees, it is important to study

the systematics of existing bee fauna and document their role in ecosystem

maintenance.

Taxonomy of Bees

Bees are insects of the superfamily Apoidea of which there are seven

major families. They are among the major groups of aculeate hymenopterans

(ants and wasps), in which most females have a modified ovipositor as a

stinging apparatus for defence or hunting or both. They differ from the other

hymenopteran insects by possessing small, distinct pronotal lobe that is well

seperated from and is below the tegula, and an extension of the pronotum

ventro-laterally as a pair of processes, one on each side that encircle or nearly

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encircle the mesosoma behind the forecoxa (Michener and Griswold, 1994;

Michener, 2000). The major families of bees include Stenotritidae, Colletidae,

Andrenidae, Halictidae, Melittidae, Megachilidae and Apidae (Michener,

2000). Several subfamilies, tribes, genera and species have been erected for

the bees of the world. Globally, over 30,000 species have so far been named,

and there are more yet to be identified.

The Tribe Anthidiini Fabricus

The Anthidiini is one of four tribes of the subfamily Megachilinae and

family Megachilidae. The others are: Megachiliini, Dioxyini and Osmiini.

This is an enormous subfamily with thousands of species that are commonly

called leaf-cutter, carder and mason bees. The anthidine bees exhibit

considerable variation in form from heriadiform, hoplitiform,

chalicodomiform, megachiliform to anthophoriform. Members of this tribe

mostly have yellow integumental markings (O’Toole and Raw, 1991;

Michener and Griswold, 1994; Michener, 2000).

According to Michener (2000), the nests of the Anthidiini are mostly

on exposed rocks, stems and leaves, or in pre-existing burrows and in their

own burrows. The entire nests (either exposed or in cavities) are constructed

from resin, sometimes mixed with or cementing together with soil particles,

pieces of leaves, chaff and plant hairs or fibers (Plate 1).

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Plate 1a, b, c, d, e, f, g and h. Anthidine bee nests and cell (s). (a) Nest of

Serapista denticulata (arrows indicating nest material made of pieces of

leaves, chaff and plant hairs). (b and c) Dorsal and anterior views of the nest

of S. denticulata (arrow indicating nest entrance). (d and e) Nests of

Afranthidium immaculatum showing nest material made of soil cemented

together with resin and plant fibers. (f, g and h) Arrows showing cells of A.

immaculatum covered with plant fibers.

a

b

c d

e

f

g h

d

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The tribe consist of thirty seven genera (see appendix 1), each with between

one and twelve subgenera.

Anthidine bees are widely distributed, occurring on all continents,

except Antarctica, and fairly common, except with only one species known

from Australia. The tribe largely comprises pollen collecting bees; the few

parasitic groups (Afrostelis, Euaspis, Hoplostelis and Larinostelis) have

narrow ranges of host taxa within the families Apidae (tribe Euglossini) and

Megachilidae (genus Megachile, Latreille) (Michener and Griswold, 1994,

Michener, 2000).

The non-parasitic genera are solitary, pollinating bees. Armbuster and

Steiner (1992) recorded two species of Pachyanthidium (Vachal) as the

principal pollinators of four species of the plants genus Dalechampia in South

Africa. Hence, it is essential to assess the phylogenetic relationship among

members of this tribe, in order to ascertain its classification status.

Justification of Research

Central to any biodiversity-dependent research is a sound knowledge

of what species are involved, particularly in pollination and pollinator studies.

This is the most basic pre-requisite for understanding broader ecological

systems as well as characterizing attributes of identified species.

However, Africa which is one of the biodiversity-rich continents in the

tropics, where a great deal of work on conservation and sustainable use of

biodiversity is urgently required is limited in resources, including taxonomic

expertise, to find out what species live within its boundary. This situation is

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not different in Ghana, where data on bee fauna seem to be non-existent. Few

of such data that exist are limited to only individual institutions within the

country. Bee pollinators like many other ecologically important animals, have

not received the needed taxonomic attention due to lack of capacity to

effectively and accurately identify existing fauna. Furthermore, virtually

nothing is known about the taxonomy and ecology of the bees that pollinate

Ghana’s food crops and wild plant species.

In the Western Palaearctic Region and South Africa, members of the

bee tribe Anthidiini have been recorded as principal pollinators of certain

plant families including the Labiatae, the Compositae, the Leguminosae and

the Zygophyllaceae (Armbuster and Steiner, 1992; Müller, 1996). In Ghana,

members of this bee tribe are known to be the principal pollinator of plants

belonging to the mint family (Labiatae) and probably many other food crops

and wild plants.

Unfortunately, many authors such as Pasteels (1984), Michener and

Griswold (1994) and Michener (2000) have questioned the generic status of

most members of the tribe. In addition, data on the phylogeny of all the

anthidine genera of the world is fragmentary and very limited to few

geographic regions. It is therefore important to study the phylogenetic

relationship among members of this tribe, in order to ascertain and probably

postulate a better classification system for the tribe.

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Objectives

Based on these taxonomic gaps in the world and Ghana in particular, the

research was principally conducted under 5 spectrums, in which the research

would seek to first and foremost, ascertain the monophyly of the Anthidiini

and evaluate the relationships among the genera as well as, revise the generic

classification of the tribe. In addition, cladistic analysis for the Anthidiini of

the world will be conducted in order to postulate their probable phylogeny or

evolutionary history. Furthermore, this research would revise classification of

the Afrotropical species of the genus Anthidiellum Cockerell and provide a

taxonomic key for the identification of the Afrotropical species of the genus

Anthidiellum.

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CHAPTER TWO

LITERATURE REVIEW

History of Classification of the Anthidiini within the Apiformes

The history of the classification of this group of closely related bees

appears in zoological records in the following chronology. An early attempt to

classify the group into higher taxa was made by Fabricus (1804). He named

the first anthidine genus, as Anthidium, and placed it in the solitary division of

the family Apiariae. Later, Lepeletier (1835, 1841) placed Anthidium

(Fabricus) with the solitary-nesting bees of the family Gastrilegides, along

with Chelostoma (Latreille), Lithurgus (Berthold) and Megachile. Schenck

(1861, 1869) subsequently placed the genus in the subfamily Megachilinae

(family Megachilidae) together with Lithurgus and Megachile. Thompson

(1872) placed Anthidium, Coelioxys (Latreille), Megachile and Stelis (Panzer)

in the tribe Megachilini (subfamily Megachilinae).

Schmiedeknecht (1882) and Friese (1895), however, used a system of

classification similar to Lepeletier (1841), placing the genus Anthidium in the

section with solitary nest-making bees of the family Megachilidae, and Stelis

with the parasitic bees in the family Stelidae. Ashmead (1899) greatly

modified Schmiedeknecht’s system, separating Anthidium from the subfamily

Megachilinae (Lithurgus, Megachile and Ctenoplectra Kirby, which is today

regarded as being in the Apidae) and placed it in the subfamily Anthidiinae of

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the family Megachilidae. He also placed Stelis and Coelioxys in separate

subfamilies, namely Stelidinae and Coelioxinae, both in the family

Megachilidae.

Robertson (1903 and 1904) in his classification of bees placed

Anthidium, Stelis and Trachusa Panzer in the family Stelidae (Anthidium and

Stelis in the subfamily Anthidiinae, and Trachusa in the subfamily

Trachusinae), both in the division Apygidialia. He based his classification on

the assumption that the pygidial plate has been lost repeatedly and

independently. In the same year, Bingham, (1903) synonymized Anthophora

gastrica (Illig), Stelis rufiventris Lepeletier, Anthidium abdominale (Smith)

and Euaspis abdominalis (Gerstaecker). E. abdominalis is presently the valid

name.

Michener (1944) provided a comprehensive classification of these

bees. He placed Anthidium, Dioxys Lepeletier and Serville, and Stelis in the

tribe Anthidiini (Megachilidae: Megachilinae). Michener and Griswold (1994)

and Michener (2000), however excluded Dioxys from the tribe Anthidiini. The

present genera of tribe Anthidiini are listed in appendix 1, and have been

grouped according to the number of female mandibular dentition.

Based on what appears to be inconsistency in the classification of the

tribe, it is vital to review the present classification system of the Anthidiini

using phylogenetic analysis.

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Overview of all the Anthidine Genera

Various authors have worked on the tribe in different geographical

regions. An account on general shape of taxa, foraging status (parasitic/non-

parasitic), history of classification, geographic distribution, as well as generic

status may be outlined as follows for each of the thirty seven genera.

Genus Acedanthidium is a slender, megachiliform, non-parasitic genus

which was first described by Michener and Griswold (1994). Later, Michener

(2000) provided brief diagnosis on this monotypic genus, which is endemic to

India.

Afranthidium is non-parasitic, small to medium sized, megachiliform

genus which was first described by Michener (1948). Later, Michener (2000)

recorded 11 subgenera in this genus; all are Afrotropical, and two of these also

occur in the Palaearctic Region.

Afrostelis is one of the anthidine bees which is endemic to Africa. This

cleptoparasitic, hoplitiform genus is monotypic. It was first described by

Cockerell (1931b). Later, Pasteels (1969, 1984), Baker (1996a), Michener and

Griswold (1994), Michener (2000) and Pauly (2001), provided diagnoses of

the genus..

The genus Anthidiellum was first described by Cockerell (1904a).

Later, Pasteels (1969) recorded six subgenera; two of these are endemic to the

Afrotropical Region, one is Afrotropical and Oriental, two are endemic to the

Orient and one is Mesoamerica and Holarctic. This genus consists of small,

robust, megachiliform to bombiform, non-parasitic bees.

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Anthidioctes--This genus was first described by Holmberg (1887,

1903). Later, Moure (1947a) redescribed an additional subgenus. It is known

that this small hoplitiform to slender megachiliform genus of pollen-collecting

bees is endemic to the Neotropical Region with a few species also occurring

in southern Africa.

Anthidioma--This non-parasitic, chalicodomiform genus is represented

by two species; namely, A. chalicodomoides Pasteels and A. murinum

Pasteels. These were first described by Pasteels (1984). Later, Michener and

Griswold (1994) and Michener (2000), provided a brief diagnosis of this

genus. The genus is endemic to southern Africa.

Fabricus (1804) first described the genus Anthidium from the type

species Apis manicata (Linnaeus). Michener (2000) listed seven subgenera.

Three of these subgenera are Palaearctic, two are endemic to Africa, one

occurs in the Nearctic Region, and the other subgenus occurs on all

continents, except Australia and it also does not occur in the Indo-Malayan

tropics. This genus comprises of small to medium size, megachiliform, pollen-

collecting bees.

Apianthidium--This megachiliform, non-parasitic genus was first

described by Pasteels (1969). It is monotypic, and occurs in Borneo.

Aspidosmia--This monotypic genus was first described by Brauns

(1926) as a subgenus of genus Osmia due to its striking morphological

similarity with species of Osmia s. str. Later, Peters (1972a) gave Aspidosmia

generic status. This genus comprises mostly of robust megachiliform, non-

parasitic bees that are endemic to southern Africa.

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Atropium--This robust megachiliform, non-parasitic genus was first

described by Smith (1853). It is known by a single female specimen of

Atropium atropos (Smith), which is endemic to Gambia.

Aztecanthidium--The genus was described first by Michener and

Ordway (1964). The only three representative species were revised by

Snelling (1987). Michener (2000) later provided a brief diagnosis on the

genus. This large-sized, chalicodomiform, pollen-collecting genus is endemic

to Mexico.

The genus Bathanthidium was first described by Mavromoustakis

(1953). Later, other authors, such as Pasteels (1968a, 1969), Romankova

(1988), and Michener and Griswold (1994) worked on the genus. Michener

(2000) listed three subgenera. Two of these subgenera are endemic to the

Oriental Region and the other occurs in the eastern Palaearctic Region. This

genus consists of small hoplotiform to slender megachiliform, non-parasitic

bees.

Pasteels (1969) described the genus Benanthis for a species that was

originally described by Benoist (1962) in the genus Anthidium. It is still the

only described species in this genus of medium to large megachiliform, non-

parasitic bees. It is endemic to Madagascar.

Cyphanthidium--This non-parasitic, medium sized, megachiliform

genus is represented by two species. It was first described by Pasteels (1969),

and it is known to comprise two representative species. Cyphanthidium is

endemic to southern Africa.

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Dianthidium--This genus was first described by Cockerell (1900).

Later, other authors such as, Moure (1965), Griswold and Michener (1988),

and Michener (1942, 2000) worked on the group. Michener (2000) recorded

four subgenera. One is endemic to the Nearctic Region; two are only known

from Mexico, and two occur in South Africa. It consists of mostly small to

medium sized, chalicodomiform, pollen-collecting bees.

Duckeanthidium--This non-parasitic, robust, chalicodomiform genus

was first described by Moure and Hurd (1960). Later, five species were

described by Urban (1995). Michener (2000) synonymized Grafanthidium and

Duckeanthidium. All of the species are endemic to Brazil.

Eoanthidium--Popov (1950) first described this non-parasitic, small

megachiliform to slender bombiform genus. Michener (2000) listed four

subgenera, two of which are endemic to Africa and the Palaearctic Regions.

The other two occur in India.

Epanthidium--Moure (1947a) first described this non-parasitic genus.

Later, Stange (1983), Moure and Urban (1990) Urban (1991) and Michener

(2000) worked on this genus. Michener (2000) recorded three subgenera, two

are endemic to South America and the other occurs in both South America

and Mexico. It comprises small chalicodomiform bees.

Euaspis--This cleptoparasitic, medium to large, chalicodomiform

genus was first described by Gerstaecker (1857). Since then, other authors,

such as Fairmaire (1858), Vachal (1903), Friese (1909a, 1912b, 1914, and

1921) and Cockerell (1933e), worked on this genus. Michener (2000)

recorded 12 species two were listed as Afrotropical. The other ten are

Oriental.

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Gnathanthidium--This non-parasitic genus was first described by

Pasteels (1969). It is monotypic and endemic to eastern and southern Africa.

Hypanthidiodes--This genus was first described by Moure (1947a).

Michener (2000) listed ten subgenera, all of which are Neotropic. This non-

parasitic genus comprises small to medium-sized chalicodomiform or

hoplitiform bees.

The genus Hypanthidium was first described by Cockerell (1904b).

Later, Urban (1994a) revised this genus. Michener (2000) listed two

subgenera, all of which are endemic to the Neotropical Region. It consists of

large chalicodomiform bees.

Hoplostelis--This cleptoparasitic genus was first described by

Dominique (1898). Other authors, such as Griswold and Michener (1988),

Moure and Urban (1994) and Michener (2000), worked on this genus.

Michener (2000) listed three subgenera, all of which are Neotropical. This

genus consists of mostly medium to large sized, hoplitiform to

chalicodomiform bees.

Icteranthidium--This genus was first described by Latreille (1809).

Later, other authors, such as Michener (1948) and Pasteels (1969, 1984)

worked on this genus. According to Michener (2000), there are about 25

species, 15 species occur in the West Palaearctic Region, two are endemic to

the Afrotropical Region and the other eight, are Oriental.

Indanthidium was first described by Michener and Griswold (1994).

This monotypic genus consists of small to medium-sized, megachiliform,

pollen-collecting bees that are endemic to southern India.

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Larinostelis--This genus was first described by Michener and

Griswold (1994). It is represented by only one described species Larinostelis

scapulata Michener and Griswold that is endemic to Kenya

The genus Neanthidium was first described by Pasteels (1969). It is

represented by one species Neanthidium octodentatum (Pasteels), which

occurs in North Africa (Algeria, Morocco).

Notanthidium--This slender, hoplitiform genus of pollen-collecting

bees was first described by Isensee (1927). Later, Moure (1947a) and Urban

(1997b) worked on the genus. Michener (2000) listed three subgenera, all of

which are endemic to South America.

Pachyanthidium was first described by Friese (1905). Since then, other

authors, such as Cockerell (1930b), Pasteels (1969) and Michener and

Griswold (1994), worked on this group. Michener (2000) recorded four

subgenera. Two are endemic to Africa, one occurs in both Africa and Arabia,

while the other occurs in both Africa and the Orient. This genus comprises

non-parasitic, robust megachiliform bees.

Paranthidium was first described by Cockerell and Cockerell (1901).

Michener (1948) described an extra two subgenera in this genus (namely

Rapanthidium and Mecanthidium), but later Michener (2000) moved

Mecanthidium to the genus Dianthidium. One of the subgenera is endemic to

Mesoamerica and the other occurs in both Mesoamerica and the Nearctic

Region.

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Pleisianthidium--This genus was first described by Cameron (1905).

Since then, other authors, such as Mavromoustakis (1951), Pasteels (1969,

1984) and Michener and Griswold (1994), have revised the group. Michener

(2000) listed four subgenera, all of which are endemic to South Africa.

Pseudoanthidium was first described by Friese (1898). Michener

(2000) recorded six subgenera, two of which are endemic to Africa, three are

known from the Palaearctic and Orient regions. The other subgenus occurs in

both Africa and Arabia.

The genus Rhodanthidium was first described by Iseense (1927).

Michener (2000) recognized three subgenera, all of which occur in the

Palaearctic Region, however, one also occurs in the Oriental Region.

Serapista--This robust, megachiliform to chalicodomiform, pollen-

collecting genus was first described by Smith (1854) as Serapis. However,

because Serapis was preoccupied under phyllum mollusca, Cockerell (1904)

renamed this genus as Serapista Cockerell. Other authors, such as

Mavromoustakis (1940) and Pasteels (1969, 1984), worked on this genus.

Pasteels (1984) recorded four species all of which are endemic to Africa.

Stelis--This cleptoparasitic genus was first described by Panzer (1806).

Since then several other authors, such as Provancher (1888), Ashmead (1898),

Robertson (1902), Noskiewicz (1961) and Michener and Griswold (1994),

revised this genus. Michener (2000) listed six subgenera. Two subgenera are

endemic to the Orient, two occur in Mesoamerica, with one of these also

occurs in the Palaearctic Region, which has an additional endemic genus. The

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last subgenus is Holarctic. Michener (2000), however, mentions that its

distribution extends into Kenya, yet Eardley and Urban (in preparation) do not

record a Kenyan species but did record one species from South Africa.

Trachusa--This parasitic genus was first described by Panzer in 1806.

Since then several other authors, such as Provancher (1888), Ashmead (1898),








Trachusoides--This non-parasitic, large megachiliform genus was first

described by Michener and Griswold (1994). This monotypic genus is

endemic to southern India.

Phylogenetic Position of Bees

History and Evolution of the Apoidea

Among the bee genera, phylogenetic procedures have been employed

to provide evidence of close relationship to the sphecoid wasps Comstock

(1942). Fossil record on bees is known to be limited. It has been said that data

has been biased towards resin-collecting bees which were occasionally

trapped in the resin and were later fossilized in amber (a hard translucent

fossil resin varying in colour from yellow to light brown). Until the latter part

21

of 2006, the oldest fossil bee known was Cretotrigona pricsa Michener and

Grimaldi, from the New Jersey amber (Fig. 1; Pl. 2 (Michener and Grimaldi,

1988; Engel, 2000; Michener, 2000)).

However, latter in the year 2006, a new oldest fossilized bee

[Melittosphex burmensis Poinar and Danforth, Melittosphecidae] was

described from the early Cretaceous Burmese amber (over 100 million years

ago (Fig. 2 & Pl. 3)). M. burmensis from the Burmese amber is 35-45 million

years older than C. prisca. This new fossil provides insights into the

morphology of the earliest bees and provides a new minimum date for the

antiquity of bees and bee-mediated pollination Poinar and Danforth (2006).

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Plate 2. Cretotrigona pricsa Michener and Grimaldi from the New Jersey

amber (after Engel, 2000)

Plate 3. Melittosphex burmensis Poinar and Danforth from the Burmese amber

(after Poinar and Danforth, 2006)

A

A

23

Figure 1. Cretotrigona pricsa (Illustration after Michener and Grimaldi 1988)

Figure 2. Melittosphex burmensis (Illustration after Poinar and Danforth,

2006).

Fig 2

Fig 2

Fig 2

Fig 2

Fig 2

Fig 2

Fig 2

Fig 2

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Danforth (2004) in his review on bee phylogeny stated that

“Additional insights into the timing of bee pollination can be gained from

mapping extant pollinator relationships onto the phylogeny of the

angiosperms (Soltis, et al., 1999, Qiu, et al., 1999; the Missouri Botanical

Garden’s Angiosperm Phylogeny web site for the most up-to-date information

on this subject). Alone, this approach is fraught with problems. First,

pollination syndromes are highly variable within higher categories and it is

difficult to assign a state unambiguously to angiosperm families or orders.

Second, there is no guarantee that plant/pollinator relationships have remained

the same from the origins of the angiosperms to the present. Nevertheless,

specialized relationships between bees and angiosperms are not likely to have

existed prior to the common ancestor of the eudicots because extant

magnoliids (monocots, Winterales, Laurales, Magnoliales, Chloranthales,

Piperales, etc.) are, for the most part, not bee pollinated (Thien, et al., 2000).

The eudicots have recently been estimated to be between 147 and 131 million

years old based on combined fossil and DNA evidence (Wikstrцm, et al.,

2001). Finally, molecular estimates of angiosperm origins push the date of the

angiosperm/gymnosperm split back to approximately the Triassic/Jurassic

boundary (210 Ma; Wolfe, et al., 1989, Laroche, et al., 1995, Goremykin, et

al., 1996, and Sanderson, 1997). Needless to say, this is considerably earlier

than the oldest fossil angiosperms (early Cretaceous, 145 Ma) and casts even

more doubt on the belief that bees necessarily arose in the mid- to late

Cretaceous”.

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http://www.entomology.cornell.edu/BeePhylogeny/bibliography.html#wolfeetal1989

http://www.entomology.cornell.edu/BeePhylogeny/bibliography.html#larocheetal1995

http://www.entomology.cornell.edu/BeePhylogeny/bibliography.html#goremykinetal1996

http://www.entomology.cornell.edu/BeePhylogeny/bibliography.html#goremykinetal1996

http://www.entomology.cornell.edu/BeePhylogeny/bibliography.html#sanderson1997

25

The few anthidine fossil bees known are Anthidium exhumatum

Cockerell, Anthidium scudderi Cockerell and “Dianthidium” terttarium

(Cockerell). These fossil bees are believed to be of late Eocene- early

Oligocene period (Engel and Perkovsky, 2006)

Overview of Phylogenetic Studies in Apoidea

Roig-Alsina and Michener (1993) conducted the first phylogenetic

analysis on all the known long-tongued bee families. Two years later, the

phylogenetic analysis on short-tongued bees was conducted by Alexander and

Michener (1995). Various authors have worked on some of the bee tribes and

families. Some of these works can be found in Rozen (1970) and Engel (2002)

on the systematic and phylogenetic relationships of the Fideline bees of the

family Megachilidae. It is also known that, Brooks (1988), Alexander (1990;

1994), Roig-Alsina (1991) and Rozen (1996) have worked on the

phylogenetic studies of the Anthophorine bees whereas Rozen and McGinley

(1974) conducted phylogenetic studies on the bee family Melittidae. The

cladistic analysis on the subfamily Meliponinae was studied by Camargo and

Menezes-Pedro (1992). More research works on the phylogenetic studies of

bees can be found in Pignata and Diniz-Filho (1995) who conducted studies

on the phylogenetic autocorrelation and evolutionary constraints in worker

body size of some neotropical stingless bees. The phylogeny of the

26

Ammobatini was studied by Eardley and Brothers (1997) whiles the molecular

phylogeny and historical biogeography of the large carpenter bees, genus

Xylocopa (Hymenoptera: Apidae) have been done by Leys, et al., (2002).

Phylogenetic Position of the Anthidiini

The phylogenetic position of the Anthidiini within the family

Megachilidae (Subfamily Megachilinae) was studied by Roig-Alsina and

Michener (1993) in their cladistic analysis of long-tongued bees based on

adult and larval characters. The tribe Lithurgini emerged as the sister group of

tribes Anthidiini, Megachilini and Osmiini. The topology was supported by

six non-homoplasious synapomorphies, including the Labrum being longer

than broad, the thickening of secondary tentorial bridge merging with the

eutentorial arm before reaching the head wall, a reduced fan-shaped posterior

sheets of tentorium and the gonostylus more or less indistinguishably fused to

the gonocoxite. Furthermore, the volsella of the male is absent or fused to the

gonocoxite. Presently the tribe Dioxyini has been included within the

subfamily Megachilinae (Michener and Griswold, 1994). The monophyly of

the Anthidiini was supported by five non-homoplasious synapomorphies. The

five synapomorphies includes the anterior surface of the labrum of female

without basal polished area, the lateral expansion of internal thickening above

the epistomal ridge reduced and less than half width of socket diameter, a

lateral carina separating exposed part of the propleuron from part hidden by

pronotum is absent and the trunk of the anterior tibial spur with low expansion

at right angles to the velum, which curves apically into the spine of omaulus.

27

In addition, the stigma of the forewing is small, about as long as broad to

nearly being absent. This analysis was based on character from two anthidine

genera Roig-Alsina and Michener (1993).

In this study, a phylogenetic concept including a hypothetical

evolutionary scenario is presented for the Anthidiini of the world based on

cladistic analysis of adult morphology.

28

CHAPTER THREE

MATERIALS AND METHODS

Sources of the Bees Examined

A total of 990 bee specimens belonging to 35 genera were obtained

from various sources from August, 2005 to January, 2008. The genera and the

total number of specimens examined (in parenthesis) are as follows: Lithurgus

(2), Megachile (2), Acedanthidium (2), Afranthidium (111), Afrostelis (29),

Anthidiellum (162), Anthidioma (2), Anthidium (63), Anthidioctes (2),

Apianthidium (1), Aspidosmia (2), Aztecanthidium (2), Bathanthidium (1),

Benanthis (1), Cyphanthidium (3), Dianthidium (16), Duckeanthidium (2),

Eoanthidium (37), Epanthidium (4), Euaspis (40), Gnathanthidium (3),

Hoplostelis (2), Hypanthidiodes (2), Hypanthidium (6), Icteranthidium (2),

Indanthidium (2), Neanthidium (2), Notanthidium (2), Pachyanthidium (49),

Paranthidium (2), Plesianthidium (245), Pseudoanthidium (140),

Rhodanthidium (7), Serapista (32), Trachusa (13). The detailed biogeographic

information, collector’s name and location of depository of each specimen are

provided under the material examined (Appendix 2). The material studied

were loaned specimens obtained from the following individuals, their

institution, cities and acronyms are given in parenthesis. These include

Barbara Dombarosky of the Transvaal Museum of South Africa (TMSA),

Pretoria, South Africa, Christina Lebean, John Ascher and Jerry Rozen of the

American Museum of Natural History (AMNH), New York, USA, Eliane

29

DeConick of the Musee Royal de l'Afrique Centrale (MRAC) Tervuren,

Belgium, Frank Kock Museum Naturkunde für Humboldt (ZMHB)

Universität, Berlin, Germany); George Else formerly of the Natural History

Museum (NHM), London, United Kingdom, Jennifer Thomas of the Snow

Entomological Museum Collection (SEMC), University of Kansas, United

States of America and Margie Cockrane of the South African Museum

(SAMC), Cape Town, South Africa. Other depositories included Mary

Gikungu of the Kenya National Museum (KNM), Nairobi, Kenya, Peter

Kwapong University of the Cape Coast Entomology Museum (UCCEM),

Cape Coast, Ghana and Ros Urban of the South African National Collection

of Insects (SANC), Pretoria, South Africa.

Preparation of Specimens

Immediately the loaned specimens were received from the various

museums, they were quarantined and curated. Quarantine procedures involved

storage of specimens in freezers at temperatures of -4ºC to -15 ºC for a

minimum period of two weeks to eliminated possible museum pests such as

mites. In certain instances, direct hand picking method was used to remove

these pests. Curating of specimens was carried out to ensure that the detailed

information on each specimen conformed to what was provided by the

literature, as well as ensuring that each specimen was accurately placed into

the appropriate taxon.

30

Examination of Specimens

The morphological examination of the specimens in this present study

was on visual observation of external morphological features. Specimens were

examined, and illustrations and measurements made using an Olympus

dissection microscope fitted with a drawing tube and a graticule. Photographic

images of the specimens including certain features that were used for their

diagnoses were captured using camera lucida. The Combined ZM computer

software was used to edit captured images. Male genitalia and hidden sterna

of some relaxed specimens were removed using a pair of forceps and

macerated in 10% KOH for 12–24 hours at room temperature, depending on

the thickness of the cuticle of each structure. After clearing the genitalia and

sterna were stored and examined in 75% glycerine.

Morphological terminologies, Abbreviations and Measurements

The terminology used in this study follows that of Michener (1944,

2000). Different morphological terms are explained in the text. Vestiture

generally refers to the relatively fine hairs and where setae are specifically

mentioned, these are thick hairs. Pubescent hair refers to branched hairs, while

setose hair refers to unbranched hairs. Small punctures, as referred to in the

descriptions, measure 0.8 mm or less in diameter, whereas large punctures are

1.0 or more mm in diameter. Terminology such as antennal scape groove was

used to describe the large fovea or depression above the antennal sockets of

the female of genus Notanthidium (Fig. 3b).

31

In the revision of the genus Anthidiellum, the terms outer and inner

carinae were used to describe one or two ridge (s) on the mandible of the

female, the outer carina originates from the mesal region of the mandible and

terminates on the outer tooth, the inner carina originates from the basal region

of the mandible and terminates in the mesal region. Illustrations of the legs

exclude the coxa and the anterior region of trochanter. The abbreviations T

and S are used for the metasomal terga and sterna, respectively (e.g. T1 and

S1 refers to the first metasomal tergum and sternum, respectively). In the

illustrations of the male genitalia, the posterior end is directed towards the top

of the page. Furthermore, all labels on both photographs and illustrations

follow that of figure 3 and 4.

Phylogenetic Studies of the tribe Anthidiini

Selection of Taxa within the Tribe

During the cladistic studies of the anthidine genera, the type species of

the genus and the type species of the nominative subgenus (for large genera)

were selected and examined. However, for some genera the type species were

not available, hence other species belonging to those genera were selected for

the analysis.Two separate cladistic studies were undertaken. The first study

encorporated 35 species representing 33 anthidine genera (ingroup, listed in

the matrix) and two outgroups namely Lithurgus pullatus (Vachal) and

Megachile semiflava (Cockerell).

In the second analysis, 17 species of the genus Anthidiellum and two

outgroup taxa were studied. These represent the Afrotropical species of the

32

genus (listed in the cladogram) and the genera chosen as outgroups were

(Trachusa aquilifilum (Strand) and Eoanthidium turnericum

(Mavromoustakis)).

Roig-Alsina and Michener (1993) postulated that these outgroups used

for the phylogenetic studies, represent the recent common ancestors and the

most recent common ancestors of the tribe Anthidiini and genus Anthidiellum.

Hence, these two outgroups revealed the probable transformation series of the

character states within the subfamily Megachilinae.

Selection and Coding of Characters and Character States

Adults of each species were examined based on 51 external characters,

and each character for which distinct states occurred in different species was

included in the matrices. A number of the characters used in the present

phylogenetic analysis of the tribe were taken or modified from Roig-Alsina

and Michener (1993). Annotated lists of the characters used in the

phylogenetic reconstructions can be found in page 31. The states of each

character within the ingroup were coded as 0 representing primitive; 1, 2 and

3 referring to successive derived states. All character states in the outgroup

were considered to be pleisomorphic. Additive binary coding was employed.

Unknown character states were coded with "?" whereas inapplicable

characters were coded with "-". All characters were weighted equally. Four

characters of the male genitalia were deactived during the analysis.

33

Characters used for the Cladistic Analyses of the Anthidiini

The following characters and character states were used in the cladistic

analysis of Anthidiini.

Head

1. Paraocular carina. (0) Present. (1) Absent.

2. Dorsal region of preoccipital area, in female. (0) Gently rounded or

angulate. (1) Carinate.

3. Lateral region of preoccipital area, in female. (0) Gently rounded. (1)

Carinate.

4. Preoccipital area. (0) Without lamella. (1) Lamellate.

5. Juxtantennal carina. (0) Absent. (1) Present.

6. Subantennal suture. (0) More or less straight. (1) Curved.

7. Distance between antennal sockets. (0) More than two antennal socket

diameters. (1) Subequal to or less than two antennal socket diameters.

8. Antennal scape, posterior surface. (0) Densely pubescent. (1) Sparsely

pubescent to impubescent

9. Antennal scape groove of female. (0) Absent. (1) Present.

10. Mediolongitudinal region of face. (0) Slightly raised. (1) Strongly

raised.

11. Ventral margin of clypeus. (0) Without projections, smooth. (1)

Unevenly margined, with projections.

12. Ventro-lateral region of female clypeus. (0) Flat. (1) Tuberculate.

13. Dorsal region of clypeus of female. (0) Flat to gently convex. (1)

34

Highly elevated.

14. Mandibular teeth of female. (0) Three teeth. (1) Four or more teeth.

15. Malar area. (0) With one lower tubercle. (1) Possess several tubercles.

16. Facial vestiture. (0) Densely pubescence. (1) Sparse to impubescence.

Mesosoma

17. Dorso-lateral angle of pronotum. (0) Well developed. (1) Absent or

weakly developed.

18. Distal region of pronotal lobe. (0) Round or gently curved. (1) Sharply

pointed.

19. Omaulus. (0) Rounded. (1) Carinate.

20. Scutum length. (0) As long as broad. (1) Elongated, twice as long as

broad.

21. Scuto-scutellar suture. (0) Similar to scuto-axilla suture, closed. (1)

More open than scuto-axilla suture, foveate.

22. Posterior margin of axilla, dorsal view. (0) Does not extend laterally

beyond lateral margin of scutum. (1) Extend beyond lateral margin of

scutum.

23. Posterior region of axilla. (0) Without spine. (1) Possess spine.

24. Scutellum, dorsal view. (0) Not overhanging and concealing

propodeum. (1) Obscures propodeum, at least medially.

25. Scutellum , lateral view. (0). Rounded posteriorly, without lamella. (1)

Lamellate posteriorly.

26. Scutellum, posterior margin, dorsal view: (0) Entire. (1) Slightly

35

notched postero-medially. (2) Deeply notched postero-medially.

27. Apex of outer hind tibia spur. (0) More or less straight. (1) Strongly

curved.

28. Shape of female hind basitarsus. (0) Slender, subequal to or slightly

more than half width of hind tibia. (1) Broader than hind tibia.

29. Length of hind basitarsus. (0) Short, subequal to or less than half

length of hind tibia. (1) Long, subequal length of hind tibia.

30. Vestiture on dorsal surface of hind basitarsus. (0) Sparse to

impubescent. (1) Dense.

31. Basitibial plate in male. (0) well defined. (1) Very small or absent.

32. Tarsal claws of female. (0) Simple. (1) Cleft, inner ramus sometimes a

tooth.

33. Apex of marginal cell position. (0) Seperated from wing margin. (1)

On wing magin.

34. Apex of marginal cell, shape. (0) Rounded. (1) Pointed.

35. Stigma. (0) Broad, twice as wide as prestigma. (1) Slender, as wide as

prestigma or absent.

36. First recurrent vein of fore wing. (0) Extends distally behind first

transverse cubital vein. (1) Meets first transverse cubital vein distally

or enters first submarginal cell.

37. Second recurrent vein of fore wing. (0) Meets second transverse

cubital vein distally or extend distally behind second transverse cubital

vein. (1) Enters second submarginal cell midway.

36

38. Second recurrent vein, in profile. (0) Enters second submarginal cell or

meet second transverse cubital vein distally. (1) Extend distally behind

second transverse cubital vein.

39. Second cubital cell of fore wing. (0) Subequal half length of first

cubital cell or less. (1) More than half or subequal length of first

cubital cell.

40. Jugal lobe. (0) Long, more than half length of vannal lobe. (1) Short,

half as long as or less than half length of vannal lobe.

41. Apex of jugal lobe: (0) Broadly rounded; (1) Narrowly rounded to

slightly pointed.

42. Vein Cu-v of hind wing. (0) Straight. (1) Slants.

43. Vein Cu-v of hind wing (length). (0) Short, much less than half as long

as second absissa of vein M+Cu. (1) Long, subequal to half length of

second absissa of vein M+Cu. (2) Longer than second absissa of

M+Cu.

Metasoma

44. Scopa. (0) Present. (1) Absent.

45. Fovea on anterior region of T1. (0) Present. (1) Absent.

46. Hairy subapical bands on postgraduli on T1-T5. (0) Present. (1)

Absent.

47. Pygidial plate on female T6. (0) Present. (1) Absent.

37

48. Sternal setae of male. (0) Absent. (1) Present.

49. Gonostylus. (0) Narrow less than half width of gonocoxite. (1)

Expanded more than half width of gonocoxite.

50. Penis valves. (0) Narrowly seperated. (1) Widely seperated.

51. Apical region of penis valves. (0) Without notch on margin. (1)

Notched

38

Figures 3a, b, c and d. Labelled integumental structures, showing the

diagnostic characters used for the analyses. (a) Dorsal view of face of genus

Anthidiellum. (b) Face of genus Notanthidium, indicating the antennal scape

groove behind antennal socket and fovea on the ventral region of clypeus of

the female (Dorsal view). (c) Lateral view, face of female Aspidosmia,

showing the protruberance of the dorsal region of clypeus. (d) Mesosoma of

Anthidiellum (Pygnanthidiellum), in dorsal view.

b

a

c

d

39

Figure 4a, b and c. Labelled integumental structures, showing some of the

characters used for the analyses (continuation). (a) Dorsal view, metasoma of

Anthidiellum orichalscopatum. (b) S5 of male Anthidiellum otavicum,

showing the male sternal comb. (c) Male genitalia of Pleisanthidium.

a

c

b

40

Characters used for the Cladistic Analyses of the Genus Anthidiellum

During the cladistic studies on the genus Anthidiellum, the following

characters and character states were used:

Head

1. Distance between antennal sockets: (0) More than two antennal socket

diameters. (1) Subequal or less than two antennal socket diameters.

2. Length of subantennal suture: (0) Subequal or less than half clypeus

length. (1) More than half clypeus length.

3. Profile of subantennal suture: (0) More or less straight. (1) Strongly

arcuate.

4. Juxtantennal carina. (0) Absent. (1) Present.

5. Projections on female clypeal margin (Number): (0).Six or more. (1)

Four or less.

6. Clypeus mid-region profile: (0) Greatly raised. (1) Slightly raised.

7. Paraoccular carina: (0) Strongly developed. (1) Weakly developed.

8. Preoccipital carina (orientation): (0) Present dorsally, or without

carina. (1) Dorsolaterally present. (2) Present laterally.

9. Mandible punctation (size). (0) Small puntures on entire dorsal

surface. (1) Small at apex, large towards base.

10. Mandibular punctation (density): (0) Sparsely dense. (1) Moderately

dense.

11. Malar area of female. (0) Two tubercles. (1) Only one is present.

41

12. Mandibular carina of female (Size). (0) Large. (1) Reduced.

13. Mandibular carina of female . (0) Present with two carinae. (1) Present

with one carina.

Mesosoma

14. Pronotum with dorsal, anterior margin: (0) Strongly convex. (1)

Gently convex.

15. Profile of pronotal lobe, with respect to distal region: (0) Pronotal

carina small mesally, becoming progressively lamellate distally. (1)

Lamellate.

16. Pronotal lamella or carina: (0) Extend little mesad from lateral margin

of scutum as laterad (1) Extend nearly as far mesad from lateral

margin of scutum as laterad.

17. Tegula (structure): (0) Narrow anteriorly, lateral margin slightly

concave. (1) Not narrowed anteriorly, with lateral margin convex.

18. Axilla (posterior margin): (0) Extend laterally beyond lateral margin of

scutum. (1) Does not extend laterally beyond lateral margin of scutum.

19. Scuto-axilla suture: (0) Similar to scuto-axilla suture, closed. (1)

Dissimilar to scuto-axilla suture, open to form shiny fovea.

20. Lateral margin of scutum (in dorsal view). (0) Does not extend

longitudinally concave. (1) Extend longitudinally concave, groove

separate extended surface with carina next to tegula.

21. Scutellum in dorsal view, with respect to shape. (0) Medially

42

emarginate. (1) Truncate.

22. Hamuli of hind wing (Number). (0) Eighteen or more. (1) Eleven and

less than eighteen. (2) Ten or less.

23. Vein cu-v of hind wing. (0) Slants. (1) Straight.

24. Second recurrent vein of forewing. (0) Meet or nearly meets second

transverse cubital vein. (1) Does not meet second transverse cubital

vein (extend 0.5mm distally).

25. Vestiture on hind basitarsus. (0) Very dense. (1) Moderate to sparsely

dense.

26. Hind basitarsus (size): (0) Broad, more than half as broad as hind tibia,

or as broad as hind tibia. (1) Slender, subequal or half as broad as hind

tibia.

Metasoma

27. Metasoma punctation (Density): (0) Sparsely dense. (1). Moderately

dense.

28. Vestiture on T1-T6 of female. (0) At least moderately dense on T6. (1)

Sparsely dense.

29. S5 of males with comb. (0) Present (1) Absent

30. S5 of males with comb or setae. (0) Medial. (1) Lateral.

31. S5 of males with lateral comb (number of setae). (0) Very long, more

than twenty setae long. (1) Short, less than twenty and more than ten

setae long. (2) Very short, less than ten setae long.

32. Male gonostyli (structure). (0) Narrowed apically. (1) Expanded

43

apically.

33. Gonostylus (Profile of margin). (0) Notched inside, or without

marginal notch. (1). Notched outside.

34. Penis valves. (0) Narrowly seperated. (1) Widely seperated.

35. Apical region of penis valves. (0) Straight. (1) Curved.

Phylogenetic Procedures

Character matrices for cladistic analyses were constructed using the

WinClada computer software, version 1.00.08 Nixon, 1999-2002). The

phylogeny was created in NONA computer software (Goloboff, 1993) using

an unconstrained heuristic search strategy having Multiple TBR+TBR (mult*

max*). The search parameters were 100,000 maximum trees to keep

(Default=100), 500 replication (Default=1), 500 starting tree per replication

(Default=1) and zero (0) random time. Common cladogram measures such as

the consistency index (CI) and the retention index (RI) were used to evaluate

the fitness of the data to the cladogram. Analysis of characters as well as

character optimization using unambiguous optimization modus was performed

with the WinClada software.

Bootstrap analysis (Felsenstein, 1985) and jackknife sampling

(Lanyon, 1985; Siddall, 1996) were used to assess evidential support for

clades of cladograms. Values were calculated based on 100 replicates with 50

random sequence additions per replicate. Presentation of trees as well as

character mapping was carried out in the WinClada.

44

The search parameters stated were derived from series of preliminary

analyses by NONA on various anthidine genera based on that were several

characters selected and coded. Few characters that were observed to have

lower constituency index and retention index were replaced with new ones the

provided high CI and RI, as well as resolved collapsed nodes on the

cladogram.

Procedure for Studies on Systematic of the Taxa

During the systematic studies of the tribe Anthidiini and the genus

Anthidiellum, lengths were measured to confirm the size variations of these

bee species as cited in literature. These were the head lengths which were

measured from dorsal region of the preoccipital area to the ventral region of

the clypeus, the scutum and scutellum lengths were measured from the antero-

mesal region of the scutum to the postero-mesal region of the scutellum. The

forewing lengths were measurements made from the basal region to the tip of

the apical region of the forewing while, body lengths were measured from the

ventral region of the clypeus to the postero-mesal region of tergites six (in

females) and seven (in males).

In addition, various attributes of vestiture on the integument were

described. These attributes included variations in vestiture colour, density, and

distribution, the position of female scopa on sternite and the presence or

absence of the male sternal comb. Furthermore, descriptive and diagnostic

characters that could be used in the identification of the genera were recorded

based on the following features. These included colour distribution on the

45

integument, number of mandibular dentition in the female, number of

projections on lower clypeal margin of the female, the presence or absence of

carinae on face, curvature of the subantennal suture and hind tibial spur as

well as aperture of scuto-scutellar suture. Other features included the

orientation and shape of the posterior region of the scutellum, the presence or

absence of arolia, the number of lateral spines on metasoma, the shape of the

apical region of the male gonostylus and width of seperation between penis

valves of the male genitalia.

In addition, under each genus, detailed information on previous works

stated in zoological records and comment have been made on the general

shape of taxa, foraging status (parasitic/non-parasitic), geographic

distribution, as well as generic status of each of the thirty three genera

examined.

46

CHAPTER FOUR

RESULTS

Cladograms and Tree topologies of the Anthidiini

The results obtained from the cladistic analysis of the data matrix

(Table 1) using the heuristic search tool yielded 191 equally most

parsimonious trees with a length (L) of 86 steps, the consistency index (CI) of

56 and the retention index (RI) of 83. Figure 5 shows a typical tree topology of

most of the 191 most parsimonious trees obtained during analysis of

characters and character states by NONA computer software. However, the

strict consensus tree in figure 11 yielded L: 93 steps, CI: 52% and RI: 80%.

On the strict consensus tree 17 unsupported nodes were collapsed into several

polytomies. In addition, bootstrap and jackknife search strategies resulted in

maximum branch length, consistency and retension indices of 90/ 89 steps, 54/

55 % and 81/ 81 %, respectively (Figures 7 and 8).

Three major clades parallel to the outgroups (Lithurgus and

Megachile) emerged from the cladograms (Figs. 6-8) as follows:

(1) Aspidosmia, Afranthidium, Anthidioma and Neanthidium.

(2) Afrostelis, Hoplostelis, Anthidioctes, Hypanthidiodes,

Cyphanthidium, Paranthidium, Eoanthidium, Epanthidium, Euaspis,

Serapista, Apianthidium, Aztecanthidium and Duckeanthidium.

47

(3) Anthidium, Dianthidium, Icteranthidium, Anthidiellum,

Bathanthidium, Gnathanthidium, Pachyanthidium, Pseudoanthidium,

Acedanthidium, Benanthis, Hypanthidium, Indanthidium, Notanthidium,

Pleisanthidium, Rhodanthidium and Trachusa.

On the cladograms in figures 5 and 6, the symbols“○”and “●”

represent the homoplasious and non-homoplasious states in which the

characters occurred after the analysis. The tree yielded 27 of the non-

homoplasious characters. The three red circles on figures 6, 7 and 8 are

representing the three major islands revealed by the cladograms.

The acronyms attached to each taxa in all cladograms give an

indication of the biogeographic data of that genus. Thus, AF represents the

Afrotropical Region (Sub-Saharan, East and West Africa and Madagascar);

PA refers to the Palaearctic Region (North Africa and Europe); OR represents

the Oriental Region (Asian countries); NE represents the Neotropical Regions

(South America) and COS refers to the Cosmopolitan regions (refers to those

genera that occur in all the geographical areas).

During the statistical analysis of the data matrix by NONA (computer

software), all the 51 characters possessing equal weights and were of additive

coding, with 308 maximum steps and 53 minimum steps of character change.

However, 47 out of the 51 external morphological characters were actually

employed (activated), while the remaining four characters on male genitalia

were deactivated. In addition, 4 characters of male genitalia of species

Anthidioma murinum resulted as inapplicable characters "-" due to the fact

that, this genus is known by only female species. 18 characters states on males

48

from five genera were revealed as missing characters. The five genera

included Icteranthidium, Bathanthidium, Gnathanthidium, Aztecanthidium and

Duckeanthidium. The 18 characters states were examined on the genitalia of

the male specimens of these genera. However, only type specimens of those

genera were available for examination hence, examinations were not

conducted on hidden structures such as the male genitalia that require

dissection procedures. In genera Bathanthidium and Gnathanthidium only

female specimens were available. During the preliminary analysis of the tribe

by NONA, the absence of these characters was observed to have caused the

collapse of several nodes into polytomies. Hence, all those characters were

deactivated during the analysis of the tribe in the course of the main research

analysis by NONA.

49

TABLE 1. Data matrix of morphological character state of the tribe

Anthidiini indicating taxa, statistics, character and character state

of each species

TAXA CHARACTERS AND CHARACTER STATES

NTAX= 35

INFORMATIVE: 255

MAXSTEPS 308:

MINSTEPS 53:

WTS 51/51:

ACTIVATION: ]4: [47:

ADDITIVE -0: +51:

NCHAR=51

7+9283++06 +001360310 442768205+ 0082+42691 0+42130+++

8++394++17 +112471421 55387+316+ 1193+537+2 1+632411++

1111111111 1111111111 1111111111 1111111111 11111111111

1111111111 1111111111 1111111111 1111111111 11111111111

yyyyyyyyyy yyyyyyyyyy yyyyyyyyyy yyyyyyyyyy yyyyyyynnnn

++++++++++ ++++++++ + ++++++++ ++++++++++ +++++++++++

Lithurgus pullatus

Megachile semiflava

Acanthidium batrae

Afranthidium abdominale

Afrostelis tegularis

Anthidiellum rubellum

Anthidioctes willineri

Anthidioma murinum

Anthidium pontis

Apianthidium apiforme

Aspidosmia arnoldi

Aztecanthidium xochipillium

Bathanthidium sibiricum

Benanthis madagascariensis

Cyphanthidium sheppardii

Dianthidium curvatum

Duckeanthidium megachiliforme

Eoanthidium rothschildi

Epanthidium trigrinum

Euaspis abdominalis

Gnathanthidium prionognathum

Hoplostelis catamercensis

Hypanthidiodes argentina

Hypanthidium flavomarginatum

Icteranthidium laterale

Indanthidium crenulaticauda

Neanthidium octodentatum

Notanthidium steloides

Pachyanthidium bicolor

Paranthidium perpictum

Pleisianthidium fulvopilosum

Pseudoanthidium truncatum

Rhodanthidium siculum

Serapista denticulata

Trachusa aquilifilum

0000000000 0000000000 0000000000 0000000000 00000000000

1000000000 0001001000 0000000000 1100001000 10000011010

0100000000 0001011010 1111000001 1100100111 11001110100

1010001000 0001001010 0001000001 1101010101 10011110100

1010001100 0001011011 0001000000 1100000101 10001110000

1110010000 1001011010 1001010001 1100000111 11001110000

1010101100 0001011010 0001000000 1100000101 10001110000

1010001000 0001001010 0001000001 1101010101 1000101------

1010000000 0001001010 0001000001 1100000011 11001110100

1010101101 1001111110 0001111000 1110101111 10101110110

1010001000 0011001010 0000000001 1100001001 10001110110

1010101101 1001111110 0001111000 1110100111 10101110???

1110010000 0001011010 1001000001 1100000111 1100111????

0100000000 0001011010 0000000001 1100100111 11001111110

1010001100 0001011010 0001000000 1110000111 10001110010

1110000000 0000011010 0001000001 1100000111 11001111000

1010101101 1001111110 0001011001 1110100111 10101110???

1010111101 1001011010 1001110000 1110000111 10001110000

1010101101 1001011010 0001110000 1110000111 10001111000

1010101101 1001011010 0001120000 1110000111 10011110100

1111000000 0001011010 0101000101 1100010111 1100111????

1010001100 0001011010 1001000000 1100000101 10011111010

1010101100 0001011010 0001000000 1100000101 10001111000

0100000000 0001011010 0000000011 1100100011 11001110111

1110000000 0001011010 0111000001 1100000111 11001110???

0100000000 0001011010 0001000011 1100110111 11001110010

1010001000 0001001010 0001000001 1101010101 10001111000

0100000010 0101011010 0000000011 1100100111 11001110010

1111000000 1001011010 0101100001 1100000111 11001111000

1010001100 1001011010 0001010000 1110000111 10001111110

0100000000 0001011010 0000000011 1100100111 11001111100

1111010000 1001011010 0101000001 1100000111 11001111000

0100000000 0001011010 0000000011 1100100111 11001111000

1010101101 1001111110 0011110000 1110100011 10101010111

0100000000 0001011010 0001000011 1100100011 11201111100

50

Lithurgus pullatus COS

Megachile semiflava COS

Acanthidium batrae OR

Afranthidium abdominale AF, PA

Afrostelis tegularis AF

Anthidiellum rubellum COS

Anthidioctes willineri NE

Anthidioma murinum AF

Anthidium pontis COS

Apianthidium apiforme OR

Aspidosmia arnoldi AF

Aztecanthidium xochipillium NE

Bathanthidium sibiricum OR

Benanthis madagascariensis AF

Cyphanthidium sheppardii AF

Dianthidium curvatum NE

Duckeanthidium megachiliforme NE

Eoanthidium rothschildi AF, OR, PA

Epanthidium trigrinum NE

Euaspis abdominalis AF, OR

Gnathanthidium prionognathum AF

Hoplostelis catamercensis NE

Hypanthidiodes argentina NE

Hypanthidium flavomarginatum NE

Icteranthidium laterale AF, PA

Indanthidium crenulaticauda OR

Neanthidium octodentatum PA

Notanthidium steloides NE

Pachyanthidium bicolor AR, PA

Paranthidium perpictum NE

Pleisianthidium fulvopilosum AF, OR, PA

Pseudoanthidium truncatum AF, OR, PA

Rhodanthidium siculum OR, PA

Serapista denticulata AF, OR

Trachusa aquilifilum COS

21

1

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Figure 5. Cladogram showing the hypothetical phylogeny of the tribe

anthidiini using the heuristic search tool.

51




































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Figure 6. Cladogram indicating the strict consensus tree of the data matrix of

the 191 most pasimonous trees

52

Figure 7. Cladogram showing the strict consensus tree of bootstrap analysis




































51

78

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53

Figure 8. Cladogram showing the strict consensus tree of jackknife analysis of

clades.




































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54

Systematics of the tribe Anthidiini

General diagnosis of the tribe

Based on the examination of 990 bee specimens, a catalogue on the

diagnostic features of the tribe Anthidiini is presented in telegraphic writing as

follows:

Length: head 1.2-3.0 mm; scutum and scutellum 1.3-5.2 mm; forewing 5.5-

11.2 mm; body 3.9-17.5 mm. Vestiture: mostly white, except greyish in genus

Anthidioma, reddish-brown to black in most parasitic genera and on tarsi of

most taxa; sparse to moderately pubescent; setosed hairs on tibia, basitarsi and

tarsi; female scopa usually on S2-S6, absent in parasitic genera; male sternal

comb on variable number of sclerites in Hypanthidioctes, Neanthidium,

Paranthidium, Rhodanthidium, Trachusa, except Anthidiellum, Benanthis,

Cyphanthidium, Epanthidium, Hoplostelis, Pachyanthdium, Pleisianthidium,

Pseudoanthidium, but absent in Afranthidium, Afrostelis, Anthidium,

Apianthidium, Aspidosmia, Dianthidium, Eoanthidium, Euaspis,

Hypanthidium, Notanthidium, Serapista. Integument: black with cream,

yellow, or reddish maculations, except in some few species with yellow to

reddish-brown, maculations absent in genera Afrostelis, Aspidosmia, Euaspis

and Pleisianthidium; integument punctations moderate to very dense, usually

small in most genera, shiny between punctures; most genera have various

degrees of carinae on paraocular area, preoccipital area, mandible, pronotal

lobe, postero-lateral regions of axilla and scutum; in genera Gnathanthidium

and Pachyanthidium carinae is lamellate; female mandible multidentate,

55

usually four or more teeth, except female of genus Dianthidium with three

teeth, male usually tridentate, except in genus Hypanthidium with mandible

bidentate, clypeus usually broad, dorsal region gently convex, except in genus

Aspidosmia with strongly elevated protruberance; subantennal suture mostly

straight, arcuate in Anthidiellum, Eoanthidium and Pseudoanthidium; omaulus

carinate except in Bathanthidium and Paranthidium; scutum broader than

long, uniquely elongate in genus Afrostelis; axilla rarely extend beyond lateral

regions of scutum, except in genera Acedanthidium, Serapista and

Icteranthidium; scutellum slightly overhungs propodeum in most taxa;

metasoma possess spines on lateral region in a few genera, S5 or S6 and

posterior margin of T6 of some females; T7 of male usually bifid or trifid;

hind tibia subequal or broader than hind basitarsus, except in genus

Gnathanthidium with hind basitarsus broader than hind tibia; hind tibial spur

straight, except in genera Apianthidium, Aztecanthidium and

Duckeranthidium; inner ramus of claw simple, except cleft in genus

Aspidosmia; arolia present in most genera, absent in Anthidium, Trachusa,

Hypanthidiodes and Apianthidium; male gonostylus generally more or less

fused to gonocoxite, forming gonobase.

Genus Acedanthidium Michener and Griswold

Acanthidium Michener and Griswold, 1994: 305. Type species: Acanthidium

batrae Michener and Griswold, 1994, by original designation;

Michener, 2000: 480.

56

Comment

This slender, megachiliform, non-parasitic genus was first described by

Michener and Griswold (1994) using the type species Acanthidium batrae.

Later, Michener (2000) provided brief diagnosis on this monotypic genus,

which is endemic to India.

Diagnosis

Length: head 1.8-1.9 mm; scutum and scutellum 1.8-2.1 mm; forewing

5.5-5.8 mm; body 6.7-7.0 mm. Vestiture: white; moderately dense on head and

mesosoma, sparse on tergites; female scopa on S2-S5; male sternal comb

absent. Integument: black, except yellow maculations on most of lower half of

face, most of paraocular area, entire preoccipital margin, dorsal margin of

tegula, antero-lateral region of scutum, posterior regions of axilla and

scutellum, most of dorsal surfaces of legs, subapical bands on lateral regions

of T1-T6 (slightly narrow on T1-T4, wide on T5-T6 (Pl. 4a)); female mandible

with four teeth; lower clypeus margin with three projections; subantennal

suture straight; juxtantennal carina absent; preoccipital area with dorsal carina;

postero-lateral region of axilla slightly extend beyond lateral magin of scutum

(Pl. 5); scuto-scutellar suture slightly open, forming narrow fovea, scuto-axilla

suture closed; scutellum, in dorsal view, rounded posteriorly, slightly obscures

propodeum; arolia present; hind tibial spur curved; lateral spines on T5, T6;

male gonostylus expanded apically; penis valves narrowly seperated, arcuate

apically (Figure 9a).

57

Figure 9a, b and c. Male genitalia of three anthidine genera. (a) Acedanthidium

batrae Michener and Griswold. (b) Afranthidium abdominale (Friese). (c)

Anthidium cordiforme (Friese).

a

b Fig. 15

c

58

Plate 4a, b and c. Anthidine bees. (a) Female of Acedanthidium batrae

Michener and Griswold. (b) Afranthidium abdominale (Friese). (c) Afrostelis

tegularis (Cockerell).

Plate 5. Axilla of A. batrae Michener and Griswold indicating it extended

lateral region.

Plate 5

a

b

c

59

Genus Afranthidium Michener

Subgenus Afranthidium Michener s. str.

Anthidium (Afranthidium) Michener, 1948: 24. Type species: Hypanthidium

haplophilum Cockerell, 1936 by original designation.

Afranthidium (Afranthidium) Michener: Pasteels, 1984: 76-78; Michener and

Griswold, 1994: 309; Michener, 2000: 482.

Subgenus Branthidium Pasteels

Branthidium Pasteels, 1969: 88. Type species: Anthidium braunsi Friese,

1904, by original designation.

Honanthidium Pasteels, 1969: 88. Type species: Anthidium honestum

Cockerell, 1936, by original designation; Pasteels, 1984: 59, syn.

Afranthidium (Branthidium) Pasteels: Michener and Griswold 1994: 309;

Michener 2000: 482.

Subgeus Capanthidium Pasteels

Capanthidium Pasteels, 1969: 85. Type species: Anthidium capicola Brauns,

1905, by original designation; Pasteels, 1984: 64.

Afranthidium (Capanthidium) Pasteels: Michener and Griswold, 1994: 310;


Subgenus Domanthidium Pasteels

Domanthidium Pasteels, 1969: 95. Type species: Anthidium abdominale

Friese, 1904, by original designation.

Afranthidium (Domanthidium) Pasteels: Michener, 2000: 483.

Subgeus Immanthidium Pasteels

Immanthidium Pasteels, 1969: 89. Type species: Anthidium immaculatum

Smith, 1854, by original designation; Pasteels, 1984: 51.

60

Afranthidium (Immanthidium) Pasteels: Michener, 2000: 483.

Subgenus Mesanthidiellum Pasteels

Mesanthidiellum Pasteels, 1969: 83. Type species: Mesanthidiellum amoenum

Pasteels, 1969, by original designation; Pasteels, 1984: 71.

Afranthidium (Mesanthidiellum) Pasteels: Michener, 2000: 483.

Subgenus Mesanthidium Pasteels

Mesanthidium Popov, 1950: 316. Type species: Anthidium pentagonum

Gussakovskiji, 1930, by original designation.

Afranthidium (Mesanthidium) Popov: Michener, 2000: 484.

Subgenus Nigranthidium Pasteels

Melanthidium Pasteels, 1969: 90 (not Cockerell 1947). Type species:

Anthidium concolor Friese, 1913, by original designation.

Nigranthidium Pasteels, 1984: 57. Replacement for Melanthidium Pasteels,

1969.

Warnckeia Pagliano and Scaramozzino, 1990: 6. Replacement for

Melanthidium Pasteels, 1969; Pasteels, 1984: 57.

Afranthidium (Nigranthidium) Pasteels: Michener and Griswold, 1994: 310;


Subgenus Oranthidium Pasteels

Oranthidium Pasteels, 1969: 95. Type species: Anthidium oraniense Brauns,

1905, by original designation; Pasteels, 1984: 83.

Afranthidium (Oranthidium) Pasteels: Michener, 2000: 484.

Subgenus Xenanthidium Pasteels

Xenanthidium Pasteels, 1984: 33. Type species: Xenanthidium biserratum

61

Pasteels, 1984, by original designation.

Afranthidium (Xenanthidium) Pasteels: Michener and Griswold, 1994: 310;


Subgenus Zosteranthidium Michener and Griswold

Afranthidium (Zosteranthidium) Michener and Griswold, 1994: 310. Type

species: Nigranthidium tergofasciatum Pasteels, 1984, by original

designation; Michener, 2000: 485.

Comment

This non-parasitic, small to medium sized, megachiliform genus was

first described by Michener (1948). Later, Michener (2000) recorded 11

subgenera in this genus; all are Afrotropical, and two of these also occur in the

Palaearctic Region. The species chosen for the cladistic analysis were

Afranthidium abdominale. This is because it is the type species of the most

diverse subgenera.

DIAGNOSIS


4.0-5.8 mm; body 5.5-8.1 mm. Vestiture: white to reddish-brown; sparse to

impubescent, except moderately dense on head and mesosoma in subgenus

Immanthidium; setose hair on tibia; scopa on female S2-S5; male sternal comb

absent. Integument: black (metasoma brown in A. folliculosum) with pale

yellow maculations on entire posterior margins of female T1-T5 and male T1-

T6 (Pl. 4b); lower clypeal margin with six projections; female mandible with 5

62

to 18 sharp teeth; subantennal suture straight; juxtantennal carina absent;

preoccipital area carinate laterally, except subgenus Branthidium with

preoccipital area rounded; scuto-scutellar and scuto-axilla sulci closed;

scutellum, in dorsal view, truncate posteriorly, sometimes obscures

propodeum; arolia absent; T1-T3 without teeth, female T4-T5 with reduced

lateral teeth, male T4-T5 variable; female T6 with posterior margin

denticulate; gonostylus usually expanded apically, inner margin slightly

notched; penis valves slender apically, mostly narrowly separated (Figure 9b).

Genus Afrostelis Cockerell

Afrostelis Cockerell, 1931b: 340. Type species: Afrostelis tegularis Cockerell,

by original designation; Michener and Griswold, 1994: 312; Baker,

1996a: 1-3; Michener, 2000: 485.

Stelis (Afrostelis) Cockerell, 1969: 57; Pasteels, 1984: 134.

Comment

This cleptoparasitic, hoplitiform genus is monotypic. It was first

described by Cockerell (1931b). Later, Pasteels (1969, 1984), Baker (1996a),

Michener and Griswold (1994), Michener (2000) and Pauly (2001), provided

diagnoses of the genus. The genus is endemic to Africa.

Diagnosis


5.5-6.2 mm; body 3.9-5.6 mm. Vestiture: white; mostly sparse to impubescent;

63

female lacks scopa; male sternal comb absent. Integument: entirely black (Pl.

4c); female mandible with four dissimilar teeth; lower clypeal margin with

three projections; subantennal suture straight; juxtantennal carina absent;

preoccipital area carinate dorsolaterally; scutum uniquely elongated (1:1.5X

length of mesosoma); scuto-scutellar suture closed, similar to scuto-axilla

suture; scutellum, in dorsal view, rounded posteriorly; arolia present;

metasoma lacks lateral teeth; gonostylus of male narrows apically, slightly

notched on outer margin; penis valves narrowly separated.

Genus Anthidiellum Cockerell

Subgenus Ananthidiellum Pasteels

Anthidiellum (Ananthidiellum) Pasteels, 1969: 49-50. Type species: Anthidium

anale Friese, 1914, by original designation; Michener and Griswold,

1994: 312; Michener, 2000: 486.

Subgenus Anthidiellum Cockerell s. str.

Anthidium (Anthidiellum) Cockerell, 1904b: 3. Type species: Trachusa

strigata Panzer, 1805, by original designation.

Anthidium (Cerianthidium) Friese, 1932a: 304. Type species: Trachusa

strigata Panzer, 1805. Isogenotypic with subgenus Anthidiellum.

Anthidiellum (Anthidiellum) Cockerell: Pasteels, 1969: 45; Pasteels, 1984: 92;

Michener and Griswold, 1994: 312; Michener, 2000: 486.

Subgenus Chloranthidiellum Mavromoustakis

Anthidiellum (Chloranthidiellum) Mavromoustakis, 1963a: 491. Type species:

Anthidium flavescens Friese, 1925, by original designation.

64

Anthidiellum (Chloranthidium) [!] Mavromoustakis: Pasteels, 1969: 48.

Anthidiellum (Chloranthidiellum) Mavromoustakis: Pasteels, 1984: 93. Type

species: Anthidiellum minutulum flavescens Friese, 1925; Pasteels,

1984: 92; Michener and Griswold, 1994: 312; Michener, 2000: 487.

Subgenus Clypanthidium Pasteels

Bathanthidium (Clypanthidium) Pasteels, 1968a: 1060; 1969: 53. Type

species: Anthidium ruficeps Friese, 1914, by original designation.

Anthidiellum (Clypanthidium) Pasteels: Michener and Griswold, 1994: 312;


Subgenus Pycnanthidium Krombein

Pycnanthidium Krombein, 1951: 292. Type species: Pycnanthidium solomonis

Krombein, 1951, by original designation.

Pygnanthidium [!] Krombein: Mavromoustakis, 1963: 491.

Pygnanthidium [!] (Pygnanthidiellum) Mavromoustakis, 1963a: 492. Type

species: Anthidium zebra Friese, 1904, by original designation.

Anthidiellum (Pycnanthidium) Krombein: Pasteels, 1969: 45; Michener and


Anthidiellum (Pygnanthidiellum) Mavromoustakis: Pasteels, 1984: 94.

Subgenus Ranthidiellum Pasteels

Anthidiellum (Ranthidiellum) Pasteels, 1969: 48. Type species:

Protoanthidium rufomaculatum Cameron, 1902, by original

designation; Michener and Griswold, 1994: 312; Michener, 2000: 487.

Anthidiellum (Rhanthidiellum) [!] Pasteels: Pasteels, 1972: 102.

65

Comment

The genus Anthidiellum was first described by Cockerell (1904a).

Later, Pasteels (1969) recorded six subgenera; two of these are endemic to the

Afrotropical Region, one is Afrotropical and Oriental, two are endemic to the

Orient and one is Mesoamerica and Holarctic. This genus consists of small,

robust, megachiliform to bombiform, non-parasitic bees.

Diagnosis


3.2-5.0 mm; body 4.0-7.0 mm. Vestiture: white to reddish-black; sparse to

impubescent; female scopa densely setose on S2-S5; male S5 with comb,

except in nominative subgenus. Integument: mostly black (Chloranthidiellum

with brown metasoma), limited yellow maculation usually on lower half of

face, lateral region of metasoma; mandible of female with four teeth; lower

clypeal margin with four or six projections; subantennal suture strongly

arcuate; juxtantennal carina absent; preoccipital carina present dorsally, except

in nominative subgenus with preoccipital area carinate dorso-laterally and in

subgenus Chloranthidiellum where carina occurs laterally only; scuto-scutellar

suture open, forms shiny fovea; axilla-scutellar suture closed; arolia present;

metasoma lacks lateral spines; gonostylus usually expanded apically, mostly

notched on outer margin; penis valves narrowly separated.

66

Genus Anthidioctes Moure

Subgenus Anthidioctes Holmberg s. str.

Anthidioctes Holmberg, 1887: 36, nomen nudum.

Anthidioctes Holmberg, 1903: 435. Type species: Anthidioctes megachiloides

Holmberg, 1903, designation of Cockerell, 1927: 2; Michener, 1948:

20

Nananthidium Moure, 1947a: 26. Type species: Nananthidium bettyae Moure,

1947, by original designation; Moure, 1947a: 26, syn.

Anthidioctes (Nananthidium) Moure: Griswold and Michener, 1988: 30;

Michener, 2000, syn.

Anthidioctes (Anthidioctes) Holmberg: Michener, 2000: 491.

Subgenus Bothranthidium Moure

Bothranthidium Moure, 1947a: 23. Type species: Bothranthidium lauroi

Moure, 1947, by original designation

Anthidioctes (Bothranthidium) Moure: Michener, 1948: 21; Michener, 2000:

491.

Comment

This genus was first described by Holmberg (1887, 1903). Later,

Moure (1947a) redescribed an additional subgenus. It is known that this small

hoplitiform to slender megachiliform genus of pollen-collecting bees is

endemic to the Neotropical Region with a few species also occuring in

southern Africa.

67

Diagnosis


3.4-3.9 mm; body 4.5-5.6 mm. Vestiture: white to yellow; sparse to

impubescent; scopa on female S2-S5; male sternal comb absent. Integument:

mostly black or yellowish brown on head, mesosoma and metasoma mostly

black, yellowish-brown pronotum, tegula and legs, yellow on anterolateral

margin of scutum, most of axilla and scutellum, lateral regions of T1-T2, sub-

apical regions of T3-T5 (Plate 6a); female mandible with four dissimilar teeth;

lower clypeal margin with four projections; subantennal suture straight;

juxtantennal carina well developed; preoccipital area carinate laterally; scuto-

scutellar suture open, forming shiny fovae, dissimilar to scutoaxilla suture;

scutellum, in dorsal view, truncate posteriorly; arolia present; metasoma lacks

lateral spines; gonostylus of male slender apically; penis valve widely

seperated.

Genus Anthidioma Pasteels

Anthidioma Pasteels, 1984: 34. Type species: Anthidioma chalicodomoides

Pasteels, 1984, by original designation.

Comment

This non-parasitic, chalicodomiform genus is represented by two

species; namely, A. Chalicodomoides Pasteels and A. Murinum Pasteels. These

were first described by Pasteels (1984). Later, Michener and Griswold (1994)

and Michener (2000), provided a brief diagnosis of this genus. The genus is

endemic to southern Africa.

68

Diagnosis

Male: Unknown. Length: head 2.1-2.4 mm; scutum and scutellum 2.6-

3.0 mm; forewing 5.5-6.0 mm; body 7.4-8.3 mm. Vestiture: grayish white,

moderately dense; postgradular margins of T2-T5 with dense hair bands;

female scopa on S2-S6. Integument: black, except yellow maculations on most

of lower half of face, entire dorsal margin of tegula, dorsolateral margins of

axilla and scutellum, fore, middle and hind tibiae and basitarsi, subapical

bands on T1-T5 (entire width on T1-T4, medial third on T5 (Plate 6b)); female

mandible with seven teeth; lower clypeus margin with four reduced

projections; subantennal suture straight; uxtantennal carina absent;

preoccipital area gently rounded; scuto-scutellar suture open, forming shiny

fovae, with scuto-axilla suture closed; scutellum, in dorsal view, rounded

posteriorly, slightly obscures propodeum; arolia absent; metasoma lacks

lateral spines; posterior margin of female T6 denticulate.

Genus Anthidium Fabricus

Subgenus Anthidium Fabricus s. str.

Anthidium Fabricus, 1804: 364. Type species: Apis manicata Linnaeus, 1758,

by designation of Latreille, 1810: 439.

Anthidium (Anthidium) Fabricus: Pasteels, 1984: 11-17; Michener and

Griswold, 1994: 314; Michener, 2000: 489-490.

Anthidium (Stenanthidium) Moure, 1947a: 16. Type species: Anthidium

espinosai Ruiz, 1938, by original designation; Michener, 1948: 16;

69

Griswold and Michener, 1988: 29, syn.

Anthidium (Tetranthidium) Moure, 1947a: 15. Type species: Anthidium latum

Schrottky, 1902, by original designation; Michener, 1948: 16;

Griswold and Michener, 1988: 29, syn.

Anthidium (Melanthidium) Cockerell, 1947: 106, not Melanthidium Pasteels,

1969. Type species: Melanthidium carri Cockerell, 1947, by original

designation; Michener, 1948: 16; Griswold and Michener, 1988: 299,

syn.

Anthidium (Melanoanthidium) Tkalců, 1967: 91. Type species: Anthidium

montanum Morawitz, 1864, by original designation; Tkalcu: 1967: 91,

syn.

Anthidium (Echinanthidium) Pasteels, 1969: 101. Type species: Anthidium

echinatum Klug, 1864, by original designation; Pasteels, 1969:101,

syn.

Anthidium (Pontanthidium) Pasteels, 1969: 105. Type species: Anthidium

pontis Cockerell, 1933, by original designation; Pasteels, 1969: 105,

syn.

Anthidium (Ardenthidium) Pasteels, 1969: 103. Type species: Anthidium

ardens Smith, 1879, by original designation; Pasteels, 1969: 103, syn.

Anthidium (Morphanthidium) Pasteels, 1969: 423. (Synonym of

Ardenthidium, because two of the included species were clearly

intended by Pasteels to be the same as Ardenthidium, Michener, 2000).

Subgenus Callanthidium Cockerell

Callanthdium Cockerell Cockerell, 1925: 365. Type species: Anthidium

70

illustre Cresson, 1879, by original designation.

Anthidium (Callanthidium) Cockerell: Michener, 2000: 490.

Subgenus Gulanthidium Pasteels

Anthidium (Gulanthidium) Pasteels, 1969: 101. Type species: Anthidium

anguliventre Morawitz, 1888, by original designation; Michener and


Subgenus Nivanthidium Pasteels

Anthidium (Nivanthidium) Pasteels, 1969: 106. Type species: Anthidium

niveocinctum Gerstaecker, 1857, by original designation; Michener

and Griswold, 1994: 313; Michener, 2000: 490.

Subgenus Proanthidium (Friese)

Anthidium (Proanthidium) Friese, 1898: 101. Type species: Anthidium

oblongatum Latreille, 1809, by designation of Cockerell, 1909: 269;

Michener and Griswold, 1994: 314; Michener, 2000: 490-491.

Subgenus Severanthidium Pasteels

Anthidium (Severanthidium) Pasteels, 1969: 106. Type species: Anthidum

severini Vachal, 1903, by original designation; Pasteels, 1984: 18-26;


Subgenus Turkanthidium Pasteels

Anthidium (Turkanthidium) Pasteels, 1969: 103. Type species: Anthidium

unicum Morawitz, 1875, by original designation; Michener and


71

Comment

Fabricus (1804) first described the genus Anthidium from the type

species Apis manicata (Linnaeus). Several authors have worked on the group

since then. The most recent author being Michener (2000) who listed seven

subgenera. Three of these subgenera are Palaearctic, two are endemic to

Africa, one occurs in the Nearctic Region, and the other subgenus occurs on

all continents, except Australia and it also does not occur in the Indo-Malayan

tropics. This genus comprises of small to medium size, megachilform, pollen-

collecting bees.

Diagnosis


4.5-7.7 mm; body 5.8-13.8 mm. Vestiture: white; sparse to moderately

pubescent, except densely setose on basitarsus; scopa on female S2-S6; male

sternal comb absent. Integument: black with extensive pallid maculation on

most of lower half of face, dorsal surface of preoccipital area, T1-T5 with

broken cross-bands, continuous in subgenus Anthidium, (maculation yellow,

except reddish in subgenera Anthidium and Severanthidium and white in

subgenus Nivanthidium); female mandible with 5-10 teeth; lower clypeal

margin with one lateral projection; subantennal suture nearly straight;

juxtantennal carina absent; preoccipital area gently rounded; scuto-scutellar

and scuto-axilla suture closed; scutellum, in dorsal view, truncate posteriorly,

sometimes obscures propodeum; arolia absent; metasomal spines variable,

72

always on male T6; female T6 without denticles; T7 with medio-longitudinal

carina, unifid (Pl. 7a); gonostylus slightly expanded apically; penis valves

widely separated (Figure 9c).

73

Plate 6a and b. Two anthidine bees. (a) Anthidioctes willineri (Moure). (b)

Anthidioma murinum Pasteels.

Plate 7a and b. Two features of anthidine bees. (a) Arrow indicating the

unified T7 of Anthidium cordiforme (Friese). (b). Arrow showing the curved

apex of hind tibial spur of Apianthidium apiforme (Meade-Waldo).

a

b

b a

74

Genus Apianthidium Pasteels

Apianthidium Pasteels, 1969: 41. Type species: Anthidium apiforme Meade-

Waldo, 1914, by original designation; Michener, 2000: 492.

Comment

This megachiliform, non-parasitic genus was first described by

Pasteels (1969) and later by Michener (2000). It is monotypic, and occurs in

Borneo.

Diagnosis

Length: head 2.6 mm; scutum and scutellum 3.2 mm; forewing 6.5

mm; body 10.3 mm. Vestiture: cream to white; sparse to impubescent; male

sternal comb absent; thick setae on tibia absent. Integument: brownish yellow,

except reddish brown maculations apex of mandible; most of supra- antennal

area, medio-longitudinal region of scutum, posterior regions of T1-T7; lower

clypeal margin with 5 projections (1 medial, 2 lateral); subantennal suture

more or less straight; juxtantennal carina absent; preoccipital area gently

rounded; scuto-scutellar suture closed; scutellum, in dorsal view, emarginate

medio-posteriorly; arolia absent; hind tibial spur curved apically (Plate 7b);

metasoma lacks lateral spines; gonostylus expand apically, notched on outer

margin; penis valves narrow apically, widely separated (Figure 10a).

75

Figure 10a, b and c. Male genitalia of three anthidine genera. (a) Apianthidium

apiforme (Meade-Waldo). (b) Aspidosmia arnoldi (Brauns). (c) Benanthis

madagascariensis (Benoist).

Pl. 1

a

2

Fig

. 17

b

Fig

. 17

c

Fig. 17

Fi

g.

19

Fig.

17

Fig

. 17

76

Genus Aspidosmia (Brauns)

Osmia (Aspidosmia) Brauns, 1926a: 208. Type species: Osmia arnoldi Brauns

1926a, monobasic.

Aspidosmia Brauns: Peters, 1972a: 167-169, 170, 171, 173, 185; Peters, 1984:

359, 362; Struck, 1992: 71, 76, 80; Struck, 1994a: 15; Michener, 2000:

492.

Comment

This monotypic genus was first described by Brauns (1926) as a subgenus of

genus Osmia due to its striking morphological similarity with species of

Osmia s. str. Later, Peters (1972a) gave Aspidosmia generic status. This genus

comprises mostly of robust megachiliform, non-parasitic bees that are endemic

to southern Africa.

Diagnosis


5.5-6.1 mm; body 8.1-10.0 mm. Vestiture: white; mostly moderately dense to

very dense; female scopa on S2-S5; male sternal comb absent. Integument:

mostly dark brown to black, yellow on male clypeus; female mandible with

three similar teeth; lower clypeal margin without projection, female clypeus,

in lateral view, uniquely tuberculate (Figure 3c); preoccipital area gently

rounded; subantennal suture straight; juxtantennal carina absent; scuto-

scutellar suture closed, similar to scuto-axilla suture; scutellum, in dorsal

view, rounded posteriorly; arolia present; inner ramus of tarsal claw uniquely

Fig.

17

77

reduced and adjacent to outer claw; metasomal without lateral teeth;

gonostylus greatly expanded apically, without marginal notch; penis valves

narrowly separated (Figure 10b).

Genus Atropium Smith

Atropium Pasteels 1984: 131-32. Type species: Megachile atropos Smith,

1853, by original designation; Michener 2000: 493.

Comment

This robust megachiliform, non-parasitic genus was first described by

Smith (1853). It is known by a single female specimen of Atropium atropos

(Smith), which is endemic to Gambia. This genus was not available for

examination during this study, due to the fact that, it is a rare specimen and

therefore, the museum was unwilling to loan it out.

Genus Aztecanthidium Michener and Ordway

Aztecanthidium Michener and Ordway, 1964: 70. Type species:

Aztecanthidium xochipillium Michener and Ordway, 1964, original

designation; Michener, 2000: 493.

Comment

The genus was described first by Michener and Ordway (1964). The only

three representative species were revised by Snelling (1987). Michener (2000)

later provided a brief diagnosis on the genus. This large-sized,

chalicodomiform, pollen-collecting genus is endemic to Mexico.

78

Diagnosis

Length: head 2.6-2.9 mm; scutum and scutellum 3.4-3.9 mm; forewing 7.7-8.1

mm; body 10.3-11.6 mm. Vestiture: cream to white; sparse to impubescent;

female scopa on S2-S6; male sternal comb absent; thick setae absent on tibiae.

Integument: mostly black with reddish or yellow maculations on most of

lower half of face of female, only on most of dorsal region of clypeus in male

(mandible black), antero-lateral region of scutum, posterior region of

scutellum, lateral regions of T1-T2, medio-posterior margin T3-T4; lower

clypeal margin with numerous projections; subantennal suture more or less

straight; marginated carina absent; preoccipital area with dorso-lateral

carina; scuto-scutellar and scuto-axilla sulci closed; posterior region of axilla

slightly produced backward (Plate 8a); scutellum, in dorsal view, emarginated

medio-posteriorly; arolia present; hind tibial spur curved apically (Plate 8b);

T5-T6 with two lateral spines per segment, T7 bilobed.

79

Plate 8a, b, c and d. Four features of the anthidine bees used for their

diagnosis. (a) Arrow showing the lateral margin of axilla of Aztecanthidium

xochipillium Michener and Ordway being similar to the margin of the scutum.

(b) Arrow indicating the curved apex of hind tibial spur of A. xochipillium

Michener and Ordway. (c) Arrows showing the bifid nature of tergite 7 of the

male Benanthis madagascariensis (Benoist). (d) Arrows showing the trifid

nature of tergite 7 of the male Cyphanthidium sheppardii (Mavromoustakis).

a

b

c d

80

Genus Bathanthidium Mavromoustakis

Subgenus Bathanthidium Mavromoustakis s. str.

Dianthdium (Bathanthidium) Mavromoustakis, 1953: 837. Type species:

Dianthidium bifoveolatum Alfken, 1937, by original designation;

Bathanthidium (Bathanthidium) Mavromoustakis: Michener and Griswold,

1994: 314; Michener, 2000: 494.

Subgenus Manthidium Pasteels

Manthidium Pasteels, 1969: 43. Type species: Anthidium binghami Friese,

1901, by original designation; Michener, 2000: 494.

Bathanthidium (Manthidium) Pasteels: Michener and Griswold, 1994: 314;


Subgenus Stenanthidiellum Pasteels

Bathanthidium (Stenanthidium) Pasteels, 1968a: 1059. Type species:

Anthidium sibiricum Eversmann, 1852, by original designation.

? Lasanthidium Romankova, 1988: 26. Type species: Stelis malaisei Popov,

1941, by original designation, syn.

Bathanthidium (Stenanthidiellum) Pasteels; Michener and Griswold; 1994:

314; Michener, 2000: 494

Comment

The genus Bathanthidium was first described by Mavromoustakis

(1953). Later, other authors, such as Pasteels (1968a, 1969), Romankova

(1988), and Michener and Griswold (1994) worked on the genus. Michener

(2000) listed

81

three subgenera. Two of these subgenera are endemic to the Oriental Region

and the other occurs in the eastern Palaearctic Region. This genus consists of

small hoplotiform to slender megachiliform, non-parasitic bees.

Diagnosis


mm; body 5.8 mm. Vestiture: white, sparse to impubescent; female scope on

S2-S6. Integument: mostly black, with limited yellow maculations paraoccular

area, lateral regions of metasoma; female mandible with four dissimilarly

teeth; lower clypeal margin without projection; subantennal suture straight;

juxtantennal carina absent; preoccipital area carinate dorsally; scuto-scutellar

and scuto-axilla suture slightly open, forming fovea; scutellum, in dorsal view,

emarginate medio-posteriorly; arolia present; metasoma lacks lateral spines.

Genus Benanthis (Pasteels)

Benanthis Pasteels, 1969: 61, 63. Type species: Anthidium madagascariensis

Benoist, 1963, by original designation; Pasteels, 1984: 133; Michener

and Griswold, 1994: 301, 314; Michener, 1997b: 10; Michener, 2000:

495; Pauly et. al., 2001: 226-228.

Comment

Pasteels (1969) described the genus Benanthis for a species that was

originally described by Benoist (1962) in the genus Anthidium. It is still the

only described species in this genus of medium to large megachiliform, non-

82

parasitic bees. It is endemis to Madagascar

Diagnosis

Female unknown. Length: head 2.0 mm; scutum and scutellum 2.1 mm;

forewing 6.1 mm; body 7.1 mm. Vestiture: white; sparse to impubescent; male

sternal comb present on S6. Integument: mostly black, tegula and legs brown,

yellow maculation on most of lower half of face, paraoccular area, dorsolateral

region of preoccital area, part of anteior region of scutum, lateral margins of

axilla and scutellum, lateral regions of T1-T6 (narrows medially on T1-T4);

lower clypeal margin without projection; subantennal suture more or less

straight; juxtantennal carina absent; preoccipital area rounded; scuto-scutellar

suture open, forming shiny fovae, scutoaxilla suture closed; scutellum, in

dorsal view, rounded posteriorly; arolia present; metasomal lateral spine

absent, T7 with unique medio-longitudinal carina, bifid (Plate 8c); gonostylus

expanded apically; penis valves widely separated, apex slightly expanded

(Figure 10c).

Genus Cyphanthidium Pasteels

Cyphanthidium Pasteels, 1969: 57. Type species: Cyphanthidium intermedium

Pasteels, 1969, by original designation; Michener and Griswold 1994:

314; Michener 2000: 495.

Trianthidiellum Pasteels, 1969: 58. Type species: Hypanthidium sheppardi

Mavromoustakis, 1937b, by original designation; Pasteels: 1969: 58,

syn.

83

Anthidiellum (Trianthidiellum) Mavromoustakis: Pasteels 1984: 113-114, syn.

Comment

This non-parasitic, medium sized, megachiliform genus is represented by two

species. It was first described by Pasteels (1969), and it is known to comprise

two representative species. Cyphanthidium is endemic to southern Africa.

Diagnosis


mm; body 8.8 mm. Vestiture: yellow to brown; sparse to impubescent; male

sternal comb absent. Integument: black, except yellow maculations on most of

lower half of face, paraoccular area, spot below medial ocellus, distal region

of pronotum, anterior margin of tegula, anterolateral margin of scutum, lateral

margin of axilla, posterolateral margin of scutellum, legs, lateral region of T1,

band on anterior surfaces of T2-T6, most of T6; female mandible with four

dissimilar teeth; lower clypeal margin with four projections; subantennal

suture more or less straight; juxtantennal carina absent; preoccipital area

gently rounded; pronotal carina uniquely lamellate; scuto-scutellar and scuto-

axilla sulci closed; scutellum, in dorsal view, truncate posteriorly; arolia

present; metasoma with lateral spines, T7 trifid (Plate 8d); gonostylus slender

apically; penis narrow apically (Figure 11a).

84

Cyphanthidium Dianthidium

Figure 11a and b. Male genitalia of two anthidine genera. (a) Cyphanthidium

sheppardii (Mavromoustakis). (b) Dianthidium curvatum (Cockerell)

a

b

85

Genus Dianthidium (Cockerell)

Subgenus Adanthidium Moure

Adanthidium Moure, 1965: 29. Type species: Anthidium texanum Cresson,

1878, by original designation

Dianthidium (Adanthidium) Moure: Michener, 2000: 496.

Subgenus Deranchanthidum Griswold and Michener

Dianthidium (Deranchanthidium) Griswold and Michener, 1988: 34. Type

species: Dianthidium chamela Griswold and Michener, 1988, by

original designation; Michener, 2000: 496.

Subgenus Dianthidium Cockerell s. str.

Anthidium (Dianthidium) Cockerell, 1900: 412. Type species: Anthidium

curvatum Smith, 1854, by original designation.

Dianthidium (Dianthidium) Cockerell: Michener, 2000: 496.

Subgenus Mecanthidium Michener

Paranthidium (Mecanthidium) Michener, 1942: 278. Type species:

Paranthidium sonorum Michener, 1942, by original designation;

Michnener, 1944: 257.

Dianthidium (Mecanthidium) Michener: Michener, 2000: 496.

Comment

This genus was first described by Cockerell (1900). Later, other

authors such as, Moure (1965), Griswold and Michener (1988), and Michener

(1942, 2000) worked on the group. Michener (2000) recorded four subgenera.

86

One is endemic to the Nearctic Region; two are only known from Mexico, and

two occur in South Africa. It consists of mostly small to medium sized,

chalicodomiform, pollen-collecting bees.

Diagnosis


4.2-7.7 mm; body 5.2-10.6 mm. Vestiture: white; sparse to impubescent;

female scopa on S2-S5; male sternal comb absent, except in Adanthidium and

Dianthidium with comb on S5; thick setae. Integument: black, except reddish

or yellow maculations on most of lower half of face, medial region of

supraclypeal area, entire paraoccular area, dorsolateral margin of preoccipital

ridge, distal margin of scutum, dorsolateral margin of axilla and scutellum,

metasoma (variable as in Plate 9a); female mandible with four dissimilar teeth;

lower clypeal margin with four reduced projections; subantennal suture more

or less straight; juxtantennal carina reduced; preoccipital ridge reduced

dorsally in most subgenera; scuto-scutellar suture closed; scutellum, in dorsal

view, truncate posteriorly; arolia present except in Deranchanthidium; lack

lateral metasomal spines; female T6 lacks carina; gonostylus narrow apically;

penis valves greatly expanded apically, narrowly separated (Figure11b, Plate

9b).

87

Plate 9a, b and c. An anthidine bee and some of the feactures used in their

diagnosis. (a) Dianthidium curvatum (Cockerell). (b) Male genitalia of D.

curvatum (Cockerell). (c). Curved apex of hind tibial spur of Duckeranthidium

theieli Michener.

b a

c

88

Genus Duckeanthidium Moure and Hurd

Duckeanthidium Moure and Hurd, 1960: 2. Type species: Anthidium

megachiliforme Ducke, 1907, by original designation; Michener, 2000:

496-497.

Grafanthidium Urban, 1995: 435. Type species: Grafanthidium amazonense

Urban, 1995, by original designation; Urban; 1995: 435, syn.

Comment

This non-parasitic, robust, chalicodomiform genus was first described

by Moure and Hurd (1960). Later, five species were described by Urban

(1995). Michener (2000) synonymized Grafanthidium and Duckeranthidium.

All of the species are endemic to Brazil.

Diagnosis


7.7-9.2 mm; body 10.5-12.3 mm. Vestiture: black; moderately dense, except

uniquely long and dense on fore, middle and hind barsitarsi and tarsi of male;

male sternal comb absent; female scopa on S2-S6. Integument: black with

reduced integumental maculation on lower region of male clypeus, lateral

regions of T1-T4; female mandible with four teeth; lower clypeal margin with

6 projections (1 medial, 2 lateral); subantennal suture more or less straight;

juxtantennal carina well developed; preoccipital area lacks carina; scuto-

scutellar suture open, forming narrow fovea; scutellum, in dorsal view,

89

emarginate medio-posteriorly; arolia present in male, absent in female; hind

tibial spur strongly curved apically (Plate 9c); T6 of male possess spine of

lateral third region, T7 with strong lateral lobe and a median spine.

Genus Eoanthidium Popov

Subgenus Clistanthidium Michener and Griswold

Eoanthidium (Clistanthidium) Michener and Griswold, 1994: 315. Type

species: Dianthidium turnericum Mavromoustakis, 1934, by original

designation; Michener and Griswold, 1994: 315; Michener, 2000: 497.

Subgenus Eoanthidium Popov s. str.

Dianthdium (Eoanthidium) Popov, 1950: 316. Type species: Anthidium

insulare, Morawitz, 1873, by original designation.

Eoanthidium (Eoanthidium) Pasteels, 1969: 51. Type species: Anthidium

elongatum Friese, 1897, by original designation; Michener and


Subgenus Hemidiellum Pasteels

Eoanthidium (Hemidiellum) Pasteels, 1972: 112. Type species: Eoanthidium

semicarinatum Pasteels, 1972, monobasic; Michener and Griswold,

1994: 315; Michener, 2000: 498.

Subgenus Salemanthidium Pasteels

Eoanthidium (Salemanthidium) Pasteels, 1969: 51. Type species:

Hypanthidium salemanse Cockerell, 1919, by original designation;


90

Comment

Popov (1950) was the first to have described this non-parasitic, small

megachiliform to slender bombiform genus. Several authors have worked on

this genus since 1950. Michener (2000) listed four subgenera, two of which

are endemic to Africa and the Palaearctic Regions. The other two occur in

India.

Diagnosis


4.8-6.3 mm; body .5.8-10.5 mm. Vestiture: yellow to brown; sparse to

impubescent; female scopa on S2-S6; male lacks sternal comb. Integument:

mostly black, Clistanthidium metasoma brownish, extensive yellow

maculation on face, dorsal margins of scutum and scutellum, entire legs,

meso-lateral regions of T1-T3 (Plates 10a & b); female mandible with four

similar teeth; lower clypeal margin with one ventro-lateral projection;

subantennal suture strongly arcuate; juxtantennal carina well developed (Plate

11a); preoccipital area gently rounded (Plate 11b); scuto-scutellar suture open,

forming shiny fovae (closed in Salemanthidium); scutellum, in dorsal view,

rounded posteriorly; arolia absent, except in Salemanthidium; T6 of male with

lateral spine; gonostylus slender apically; penis valves narrow apically (Figure

12a).

91

Figure 12a, b and c. Male genitalia of three anthidine genera. (a) Eoanthidium

rothschildi (Vachal). (b) Epanthidium trigrinum (Schrottky). (c) Euaspis

abdominalis (Fabricus)

a

c b

92

Plate 10a, b and c. Three anthidine bees, in dorsolateral view. (a) Eoanthidium

rothschildi (Vachal). (b) Eoanthidium turnericum (Mavromoustakis). (c)

Euaspis abdominalis (Fabricus).

Plate 11a, b, c and d. Four diagnostic features of anthidine bees. (a) Face of

Eoanthidium rothschildi (Vachal) with arrows indicating the juxtantennal

carina and actuate nature of the subantennal suture. (b) Postero-lateral region

of E. rothschildi (Vachal) with arrow showing the gently rounded preoccipital

area. (c) Arrow showing the straight apex of the hind tibial spur of Euaspis

abdominalis (Fabricus). (d) Arrows showing the trifid nature of tergite 7 of the

male of Euaspis abdominalis (Fabricus).

a b

a

b

c d

c

93

Genus Epanthidium Moure

Subgenus Ananthidium Urban

Ananthidium Urban, 1991: 73. Type species: Anthidium inerme Friese, 1908,

by original designation.

Epanthidium (Ananthidium) Urban: Michener, 2000: 498.

Subgenus Carloticola Moure and Urban

Carloticola Moure and Urban, 1990: 90. Type species: Dianthidium

paraguayense Schrottky, 1908, by original designation.

Epanthidium (Carloticola) Moure and Urban: Michener, 2000: 499.

Subgenus Epanthidium Moure s. str.

Epanthidium Moure, 1947a: 33. Type species: Hypanthdium tigrinum

Schrottky, 1905, by original designation; Stange, 1983: 281: 297.

Epanthidium (Epanthidium) Moure: Michener, 2000: 499.

Comment

Moure (1947a) first described this non-parasitic genus. Later, Stange

(1983), Moure and Urban (1990) Urban (1991) and Michener (2000) worked

on this genus. Michener (2000) recorded three subgenera, two are endemic to

South America and the other occurs in both South America and Mexico. It

comprises small chalicodomifom bees.

Diagnosis


4.9-6.8 mm; body 6.5-9.0 mm. Vestiture: white; moderate to sparse, except

dense on tibiae; female scopa on S2-S5; male sternal comb on S3.

94

Integument: black; limited yellow maculation on face, dorsal margin of

preoccipital rigde, most of tegula, spots on dorsal surfaces of scutum, axilla,

scutellum and metasoma; mandible of female with five dissimilar teeth; lower

clypeal margin without projection; subantennal suture straight; juxtantennal

carina well developed; preoccipital area gently rounded; scuto-scutellar scuto-

axilla suture closed; scutellum, in dorsal view, rounded posteriorly; arolia

present; metasomal lateral spines absent, except in nominative subgenus;

female with unique medial carina on T6; female S6 with unique submarginal

carina, sometimes multidentated; S7 of male trifid (subgenus Epanthidium) or

bifid (subgenera Ananthidium and Carloticola); gonostylus of male slightly

narrow apically; penis valves narrows at apex, widely separated (Figure 12b).

Genus Euaspis Gerstaecker

Euaspis Gerstaecker, 1857: 460. Type species: Thynnus abdominalis

Fabricius, 1793, by original designation; Vachal, 1903: 376; Bingham,

1903: 60; Friese, 1909a: 88, 100, 101, 325, 343, 440, 442; Strand,

1912a: 129; Strand, 1912b: 311; Strand, 1914: 67; Friese, 1914: 294;

Friese, 1921: 1104; Cockerell, 1933d: 267-268; Cockerell, 1933e: 1;

Pasteels, 1969: 64; Gess, 1981: 26, 61; Medler, 1980: 482; li Enkulu,

1988: 212; Michener, 1997: 23; Pauly, 1998: 28.

Dilobopeltis Fairmaire, 1858: 266. Type species: Diolobopeltis fuscipennis

Fairmaire, 1858 (= Anthidium abdominale Fabricius 1793), by original

designation, Thomson, 1858: 266, syn.

Parevaspis Ritsema, 1874: 1xxi. Type species: Parevaspis basalis Ritsema,

95

1874, designated by Sandhouse, 1943: 585; Baker, 1995: 282, syn.

Comment

This cleptoparasitic, medium to large, chalicodomiform genus was first

described by Gerstaecker (1857). Since then, other authors, such as Fairmaire

(1858), Vachal (1903), Friese (1909a, 1912b, 1914, and 1921) and Cockerell

(1933e), worked on this genus. Michener (2000) recorded 12 species two were

listed as Afrotropical. The other ten are Oriental.

Diagnosis


7.2-10.3 mm; body 7.5-15.7 mm. Vestiture: reddish brown to black; sparse to

impubescent; female scopa absent; male sternal comb absent. Integument:

head and mesosoma black, metasoma reddish-brown, yellow maculation

absent (Plate 10c); female mandible with four dissimilar teeth; lower clypeal

margin with six projections; subantennal suture striaght; juxtantennal carina

strongly developed, uniquely extending a little behind antennal socket;

mediolongitudinal margin of face strongly carinate; preoccipital carina

strongly developed laterally; scuto-scutellar and scuto-axilla suture closed;

scutellum, in dorsal view, emarginate posteromedially, with two flat, plate-like

lobes projecting backwards; hind tibial spur straight (Plate 11c); arolia present;

metasomal spines absent; T7 trilobed (Plate 11d); male gonostylus notched on

inner margin, apical region greatly expanded; penis valves reduced, narrowly

separated (Figure 12c).

96

Genus Gnathanthidium Pasteels

Gnathanthidium Pasteels, 1969: 92-93. Type species: Pachyanthidium

prionognathum Mavromoustakis, 1935, original designation; Pasteels

1984: 36; Michener 2000: 499-500.

Comment

This non-parasitic genus was first described by Pasteels (1969). It is

monotypic and endemic to eastern and southern Africa.

Diagnosis


6.7-7.2 mm; body 7.7-9.4 mm. Vestiture: white; sparse to impubescent; female

scopa on S2-S5. Integument: black, with reddish-brown legs; yellow or

reddish maculations on dorsolateral portion of preoccipital ridge (Plate 12c);

lateral regions of T1-T5 (Plate 12a); female mandible with 13 dissimilar teeth

(Plate 12b); lower clypeal margin with six projections; subantennal suture

straight; juxtantennal carina reduced; preoccipital area carinate or and

lamellate dorsolaterally (Plate 12c); scuto-scutellar suture closed, similar to

scutoaxilla suture; scutellum, in dorsal view, strongly truncate; arolia present;

hind barsitarsus unusually broad (Plate 12d); metasomal lateral spines absent.

97

Plate 12a, b, c and d. An anthidine bee and some of the feactures used for

diagnosis. (a) Female Gnathanthidium prionognathum (Mavromoustakis). (b)

Lower half of the face of Gnathanthidium prionognathum (Mavromoustakis)

with the arrow indicating the numerous mandibular teeth. (c) Dorsal view head

of G. prionognathum (Mavromoustakis) with the arrow showing the carinate

and lamellate nature of the preoccipital area. (d) Arrow indicating the enlarged

hind barsitarsus of G. prionognathum (Mavromoustakis).

b

d

c

a

98

Genus Hoplostelis Dominique

Subgenus Austrostelis Michener and Griswold

Hoplostelis (Austrostelis) Michener and Griswold, 1994: 676. Type species:

Stelis aliena Cockerell, 1919, by original designation; Michener, 2000:

500.

Subgenus Hoplostelis Dominique s. str.

Hoplostelis Dominique, 1898: 60. Type species: Stelis abnormis Friese, 1925,

designated by Griswold and Michener, 1988: 36.

Odontostelis Cockerell, 1931c: 542. Type species: Stelis abnormis Friese,

1925. Isogenotypic with Hoplostelis.

Hoplostelis (Hoplostelis) Dominique: Griswold and Michener, 1988: 36;

Michener, 2000: 500-501.

Subgenus Rhynostelis Moure and Urban

Rhynostelis Moure and Urban, 1994: 297. Type species: Anthidium

multiplicatum Smith, 1879, by original designation.

Hoplostelis (Rhynostelis) Urban: Michener, 2000: 501.

Comment

This cleptoparasitic genus was first described by Dominique (1898).

Other authors, such as Griswold and Michener (1988), Moure and Urban

(1994) and Michener (2000), worked on this genus. Michener (2000) listed

three subgenera, all of which are Neotropical. This genus consists of mostly

medium to large sized, hoplitiform to chalicodomiform bees.

b

c

99

Diagnosis


2.9-5.2 mm; body 4.8-7.1 mm. Vestiture: white, sparse to impubescent; female

lacks scopa; male sternal comb on S3. Integument: black, except for limited

yellow to reddish maculation on preoccipital area, legs, lateral regions of

female T1-T5, male T1-T6; female mandible usually with three dissimilar

teeth; lower clypeal margin with four small projections; subantennal suture

straight; juxtantennal carina well developed; preoccipital area slightly carinate

dorsally; scuto-scutellar suture open, forming shiny fovea; scuto-axilla suture

closed; scutellum, in dorsal view, truncate posteriorly; arolia present; S5-S6

with lateral teeth; male T7 uniquely bilobed; gonostylus slender apically; penis

valves widely seperated (Figure 13a).

GENUS Hypathidiodes MOURE

Subgenus Anthidulum Michener

Dianthidium (Anthidulum) Michener, 1948: 19. Type species: Dianthidium

currani Schwarz, 1933, by original designation.

Hypanthidiodes (Anthidulum) Michener: Griswold and Michener, 1988: 31;


Subgenus Ctenanthidium Urban

Ctenanthidium Urban, 1993a: 85. Type species: Ctenanathidium gracile

Urban, 1991, by original designation.

Hypanthidiodes (Ctenanthidium) Urban: Michener, 2000: 502.

100

Subgenus Dichanthidium Moure

Dichanthidium Moure, 1947a: 30; [also described as new later that year by

Moure, 1947b: 235]. Type species: Dichanthidium exile Moure, by

original designation.

Hypanthidiodes (Dichanthidium) Moure: Michener, 2000: 502.

Subgenus Dicranthidium Moure

Dicranthidium Moure and Urban, 1975: 837. Type species: Anthidium

arenarium Ducke, 1907, by original designation.

Hypanthidiodes (Dicranthidium) Moure and Urban: Michener, 2000: 502.

Subgenus Hypanthidiodes Moure s. str.

Hypanthidiodes Moure, 1947a: 35. Type species: Anthidium flavofasciatum

Schrottky, 1902, by original designation.

Hypanthidioides [!] Moure: Michener, 1948: 22; Moure and Urban, 1975: 837;

Urban, 1993b: 28.

Hypanthidiodes (Hypanthidiodes) Moure: Michener, 2000: 502-503.

Subgenus Larocanthidium Urban

Larocanthidium Urban, 1997a: 299. Type species: Larocanthidium

emarginatum Urban, 1997, by original designation.

Hypanthidiodes (Larocanthidium) Urban: Michener, 2000: 503.

Subgenus Michanthidium Urban

Gnathanthidium Urban, 1994a: 337 (not Pasteels, 1969). Type species:

Gnathanthidium sakagamii Urban, 1994a, by original designation.

Michanthidium Urban, 1994a: 281. Replacement for Gnathanthidium Urban,

1994a.

Hypanthidiodes (Michanthidium) Urban: Michener, 2000: 503.

101

Subgenus Mielkeanthidium Urban

Mielkeanthidium Urban, 1996: 121. Type species: Mielkeanthidium nigripes


Hypanthidiodes (Mielkeanthidium) Urban: Michener, 2000: 503.

Subgenus Moureanthidium Urban

Moureanthidium Urban, 1995: 37. Type species: Dianthidium subarenarium

Schwarz, 1933, by original designation.

Hypanthidiodes (Moureanthidium) Urban: Michener, 2000: 503.

Subgenus Saranthidium Moure and Hurd

Saranthidium Moure and Hurd, 1960: 6. Type species: Anthidium flavopictum

Smith, by original designation; Griswold and Michener, 1988: 31-32.

Hypanthidiodes (Saranthidium) Moure and Hurd, 1960: 6; Michener, 2000:

503-504.

Comment

This genus was first described by Moure (1947a). Michener (2000) listed

ten subgenera, all of which are Neotropic. This non-parasitic genus comprises

small to medium-sized chalicodomiform or hoplitiform bees.

Diagnosis


2.9-5.5 mm; body 4.8-7.6 mm. Vestiture: white; sparse dense to impubescent;

female scopa on S2-S5; male sternal comb on S3, S4, or S5. Integument:

black, yellow maculations on most of lower half of face, sometimes a spot on

dorsal

102

region of preoccipital area; anterior margin of scutum, posterior margin of

scutellum, extensively on legs and metasoma; female mandible with four

dissimilar teeth; lower clypeal margin with four reduced projections;

subantennal suture straight; juxtantennal carina well developed; preoccipital

area gently rounded; scuto-scutellar suture sightly, forming reduced fovea;

scuto-axilla suture closed; scutellum, in dorsal view, rounded or lamellate

posteriorly; arolia absent, except in subgenera Ctenanthidium and

Dichanthidium; metasomal spines absent; gonostylus of male very slender,

lacks notch on margin; penis valves slender, widely separated.

Genus Hypanthidium Cockerell

Subgenus Hypanthidium Cockerell

Hypanthidium Cockerell, 1904b: 292. Type species: Anthidium

flavomarginatum Smith, 1879, by original designation.

Hypanthdium (Hypanthidium) Cockerell: Michener, 2000: 504.

Subgenus Tylanthidium Urban

Tylanthidium Urban, 1994a: 277. Type species: Tylanthidium tuberigaster


Hypanthidium (Tylanthidium) Urban: Michener, 2000: 504.

Comment

The genus Hypanthidium was first described by Cockerell (1904b). Later,

Urban (1994a) revised this genus. Michener (2000) listed two subgenera, all

of which are endemic to the Neotropical Region. It consists of large

chalicodomiform bees.

103

Diagnosis


4.2-7.7 mm; body 9.7-12.9 mm. Vestiture: white to brownish; moderate to

sparse; female scopa present on S2-S5; male sternal comb absent. Integument:

mostly black, legs yellow to brownish; yellow maculations on most of lower

half of face; dorsal portion of preoccipital ridge; lateral margin of scutum,

axilla, posterolateral half of scutellum; most of anteromedial half of female

T1-T5, and male T6 (mediolongitudinal margin black (Plate 13a)); female

mandible with four blunt teeth, male mandible bidentate; lower clypeal margin

without projection; subantennal suture straight; juxtantennal carina absent;

preoccipital area gently rounded; scuto-scutellar suture closed, similar to

scuto-axilla suture; scutellum, in dorsal view, subtruncate posteriorly; arolia

present; T6 with lateral spines, female; male T7 trifid (Plate 14b); male

gonostylus slight expanded apically; penis valves with apex deeply notched,

widely separated (Figure 13b).

104

Hypanthidium

Figure 13a and b. Male genitalia of two anthidine bees. (a) Hoplostelis

catamercensis (Schrottky). (b) Hypanthidium flavomarginatum Smith.

Fig.

25

b

a

105

Plate 13a, b and c. Three species of the tribe Anthidiini. (a) Hypanthidium

flavomarginatum Smith. (b) Female Indanthidium crenulaticauda Michener

and Griswold. (c) Female Notanthidium steloides (Spinola).

Plate 14a, b and c. Three diagnostic features used for diagnosis of the genera.

(a) Arrow showing the lateral margin of the axilla of Icteranthidium laterale

Latreille extending beyond scutum margin. (b) Arrows showing the trifid

nature of tergite 7 of the male of Hypanthidium flavomarginatum Smith. (c)

Arrows indicating the quadrifid nature of tergite 7 of the male of Neanthidium

octodentatum (Pérez).

a

b

c

c

b a

106

Genus Icteranthidium Latreille

Icteranthidium Michener, 1948: 25. Type species: Anthidium limbiferum

Morawitz, 1875, by original designation; Pasteels 1969: 67-69; 1984:

136, 156; Michener, 2000: 504.

Comment

This genus was first described by Latrielle (1809). Later, other authors,

such as Michener (1948) and Pasteels (1969, 1984) worked on this genus.

According to Michener (2000), there are about 25 species, 15 species occur in

the West Palaearctic Region, two are endemic to the Afrotropical Region and

the other eight, are Oriental.

Diagnosis


7.4-9.9 mm; body 11.0-12.0 mm. Vestiture: white; pubescence moderate to

sparsely dense; female scopa on S2-S6; male sternal comb absent; thick setae

on tibia absent. Integument: black, except yellow maculations on lower half of

face (female mandible black), lateral region of preoccipital area, most of

dorsal surfaces of tegula and legs, lateral regions of T1-T6 (narrow on T1-T2);

mandible of female with four teeth; lower clypeal margin with 5 projections (1

medial, 2 lateral); subantennal suture more or less straight; juxtantennal carina

absent; preoccipital area lacks carina; scuto-scutellar suture closed; posterior

margin of axilla with strong spine (Plate 14a); scutellum, in dorsal view,

subtruncate posteriorly; arolia absent; T6 and T7 with reduced spines; T7

trifid.

107

Genus Indanthidium Michener and Griswold

Indanthidium Michener and Griswold, 1994: 315. Type species: Indanthidium

crenulaticauda Michener and Griswold, 1994, by original designation;

Michener, 2000: 504-505.

Comment

Genus Indanthidium was first described by Michener and Griswold

(1994). This monotypic genus consists of small to medium-sized,

megachiliform, pollen-collecting bees that are endemic to southern India.

Diagnosis


3.9-5.0 mm; body 5.1-7.1 mm. Vestiture: white; sparse to impubescent; female

scopa on S2-S6; male sternal comb absent; thick setae on tibia absent.

Integument: black, except yellow maculations on lower half of face (mandible

black), spot on dorsal region of preoccipital area, anterior margin of scutum,

dorso-lateral regions of axilla and scutellum, lateral and medial third regions

od T1-T5, medial region of T6 (Plate 13b); female mandible with seven teeth;

lower clypeal margin without projection; subantennal suture straight;

juxtantennal carina absent; preoccipital area with lateral carina; scuto-scutellar

suture closed; scutellum, in dorsal view, emarginate medio-posteriorly; arolia

present; hind tibial spur straight apically; T7 of male trifid and uniquely broad,

subequal T6; gonostylus slender apically; penis valves narrow apically, widely

separated (Figure 14a).

108

Notanthidium

Figure 14a, b and c. Male genitalia of three anthidine genera. (a) Indanthidium

crenulaticauda Michener and Griswold. (b) Neanthidium octodentatum

(Pérez). (c) Notanthidium steloides (Spinola).

a b

c

109

Genus Larinostelis Michener and Griswold

Larinostelis scapulata Michener and Griswold, 1994: 317. Type species:

Larinostelis scapulata Michener and Griswold, 1994, by original

designation; Michener 1997: 32; Michener 2000: 505.

Comment

This genus was first described by Michener and Griswold (1994). It is

represented by only one described species Larinostelis scapulata Michener

and Griswold that is endemic to Kenya. This genus was not available for

examination during this study, due to the fact that, it is a rare specimen and the

museum was unwilling to loan then out.

Genus Neanthidium Pasteels

Neanthidium Pasteels, 1969: 93. Type species: Anthidium octodentatum Pérez,

1895, by original designation; Michener, 2000: 505.

Comment

The genus Neanthidium was first described by Pasteels (1969). It is

represented by one species Neanthidium octodentatum (Pasteels), which

occurs in North Africa (Algeria, Morocco).

Diagnosis

Length: head 2.6-2.9 mm; scutum and scutellum 2.3-2.9 mm; forewing 4.8-7.1

mm; body 7.1-10.6 mm. Vestiture: white to yellow; moderate to sparse;

female scopa on S2-S5; male sternal comb on S4 and S5. Integument: black,

110

yellow or reddish-yellow maculation on most of lower half of face, dorsal

margin of preoccipital area, most of tegula, spots on dorsal surfaces of axilla

and scutellum, lateral regions of T1-T5 of female, T1-T2 of male; mandible of

female with five dissimilar teeth; lower clypeal margin with 3-5 projections, a

median and one-two lateral; subantennal suture straight; juxtantennal carina

absent; preoccipital area gently rounded; scuto-scutellar and scuto-axilla

suture closed; scutellum, in dorsal view, rounded posteriorly; arolia absent;

lateral region of T6 of female with small metasomal spines, T6 and T7 of male

with strong spines on posterior regions, multidentated (Plate 14c); S7 of male

trifid; gonostylus of male slender apically; penis valves narrows at apex,

narrowly separated (Figure 14b).

Genus Notanthidium (Isensee)

Subgenus Allanthidium Moure

Allanthidium Moure, 1947a: 21. Type species: Anthidium rodolfi Ruiz, 1938,

by original desination.

Trichanthidium Moure, 1947b: 209 (not Cockerell, 1930). Type species:

Anthidium subpetiolatum Schrottky, 1910, by original designation,

Moure, 1947a: 21, syn.

Notanthidium (Allanthidium) Moure: Michener, 2000: 505-506.

Subgenus Chrisanthidium Urban

Chrisanthidium Urban, 1997b: 181. Type species: Anthidium bidentatum

Friese, 1908, by original designation.

Notanthidium (Chrisanthidium) Moure: Michener, 2000: 506.

111

Subgenus Notanthidium Isensee s. str.

Dianthidium (Notanthidium) Isensee, 1927: 373. Type species: Anthidium

steloides Spinola, 1851, by original designation.

Notanthidium (Notanthidium) Moure: Michener, 2000: 506.

Comment

This slender, hoplitiform genus of pollen-collecting bees was first

described by Isensee (1927). Later, Moure (1947a) and Urban (1997b) worked

on the genus. Michener (2000) listed three subgenera, all of which are

endemic to South America.

Diagnosis


5.8-6.1 mm; body 7.9-10.2 mm. Vestiture: reddish brown to black; sparse

dense to impubescent; female scopa on S2-S6; male sternal comb absent.

Integument: mostly black, antenna, lateral region of preoccipital ridge, tegula,

legs reddish-brown, yellow maculations on lateral regions of T1-T3, medial

region of T4 (Plate 13c); female mandible with five teeth; lower clypeal

margin with large groove, lower margin without projection; subantennal suture

straight; juxtantennal carina absent; female with unique, distinct, antennal

scape groove, (Fig. 4); preoccipital area carinate dorsally; scutellum, in dorsal

view, rounded posteriorly; arolia present; metasomal lateral spines absent;

gonostylus slender, apically narrow; penis valve widely separated (Figure

14c).

112

Genus Pachyanthidium Vachal

Subgenus Ausanthidium Pasteels

Ausanthidium Pasteels 1969: 60, 132-133. Type species: Anthidiellum ausense

Mavromoustakis, 1934; Pasteels 1984: 130, by original designation.

Pachyanthidium (Ausanthidium) Pasteels: Michener and Griswold, 1994: 319;


Subgenus Pachyanthidium Friese s. str.

Anthidium (Pachyanthidium) Friese, 1905b: 66-67, 69; Friese, 1910: 158.

Type species: Anthidium bicolor Lepeletier, 1841, designated by

Cockerell, 1920: 298.

Pachyanthidium (Pachyanthidium): Mavromoustakis, 1963: 490; Pasteels

1968b: 482-483; Pasteels, 1984: 138-139, 141-145, 147-148, Michener

and Griswold, 1994: 319; Michener, 2000: 507.

Subgenus Trichanthidiodes Michener and Griswold

Pachyanthidium (Trichanthidiodes) Michener and Griswold, 1994: 319. Type

species: Pachyanthidium semilutum Pasteels, 1981, by original

designation; Michener 2000: 507.

Subgenus Trichanthidium Cockerell

Pachyanthidium (Trichanthidium) Cockerell, 1930b: 52. Type species:

Pachyanthidium uxtanten Cockerell, 1930, monobasic; Pasteels,

1984: 151; Michener and Griswold, 1994: 317; Michener, 2000: 507.

Comment

Genus Pachyanthidium was first described by Friese (1905). Since

113

then, other authors, such as Cockerell (1930b), Pasteels (1969) and Michener

and Griswold (1994), worked on this group. Michener (2000) recorded four

subgenera. Two are endemic to Africa, one occurs in both Africa and Arabia,

while the other occurs in both Africa and the Orient. This genus comprises

non-parasitic, robust megachiliform bees.

Diagnosis


4.3-5.8 mm; body 5.4-9.0 mm. Vestiture: white, sparse to impubescent; female

scope on S2-S6; male sternal comb on S5. Integument: mostly black, with

limited yellow maculations on most of lower half of face (Plate 16a), posterior

margin of preoccipital area, laterally on scutum, axilla and scutellum,

metasoma (Plate 15); female mandible 5 to 10 dissimilar teeth; lower clypeal

margin without projection; subantennal suture straight; suxtantennal carina

reduced; preoccipital area carinate or lamelate dorsolaterally; scuto-scutellar

and scuto-axilla suture closed; scutellum, in dorsal view, truncate (Plate 16d);

arolia absent, except in male of subgenus Trichanthidium and both sexes of

subgenus Ausanthidium; T1-T6 with lateral spines, T7 of male trifid;

gonostylus apically expanded; penis valves narrowly separated (Figure 15a).

114

Plate 15. Female of Pachyanthidium bicolor (Lepeletier).

Plate 16a, b, c and d. Four diagnostic features of two anthidine genera.

(a) Face of P. bicolor (Lepeletier), with arrows indicating the straight

nature of the subantennal suture and regions of yellow integumental

maculations. (b and c) Lower region of the face of Paranthidium

perpictum (Cockerell), arrows indicating the mandibular dentition,

projections on lower clypeal margin and subantennal suture. (d) Arrow

indicating lamella nature of the scutellum of P. bicolor (Lepeletier).

a

c

b

d

115

Pachyanthidium

Figure 15a, b and c. Male genitalia of three anthidine bees. (a)

Pachyanthidium bicolor (Lepeletier). (b) Paranthidium perpictum (Cockerell).

(c) Pleisianthidium fulvopilosum (Cameron).

a

c

b

116

Genus Paranthidium Cockerell and Cockerell

Subgenus Paranthidium Cockerell and Cockerell s. str.

Paranthidium Cockerell and Cockerell, 1901: 50. Type species: Anthidium

perpictum Cockerell, 1898, by original designation.

Paranthidium (Paranthidium) Cockerell and Cockerell: Michener, 2000: 507.

Subgenus Rapanthidium Michener

Paranthidium (Rapanthidium) Michener, 1948: 11. Type species: Anthidium

vespoides Friese, 1925, by original designation; Michener, 2000: 507.

Comment

Genus Paranthidium was first described by Cockerell and Cockerell

(1901). Michener (1948) described an extra two subgenera in this genus

(namely Rapanthidium and Mecanthidium), but later Michener (2000) moved

Mecanthidium to the genus Dianthidium. One of the subgenera is endemic to

Mesoamerica and the other occurs in both Mesoamerica and the Nearctic

Region.

Diagnosis


6.1-7.1 mm; body 7.1-9.0 mm. Vestiture: yellowish to white; sparse to

impubescent; female scopa on S2-S6; male sternal comb on S4-S5.

Integument: black, except for yellow maculation on face, dorsal margin of

preoccipital area, entire margin of scutum, axilla and scutellum, spots on

dorsal surfaces of femora and tibiae, lateral regions of T1-T3 (shorter and

117

wider on T3), most of T4-T5 (mediolongitudinal region black), entire T6-T7;

female mandible with four dissmilar teeth; lower clypeal margin with six

projections (Plate 16b and c); subantennal suture straight; juxtantennal carina

absent; preoccipital area rounded; scuto-scutellar suture closed, similar to

scuto-axilla suture; scutellum, in dorsal view, medially emarginate, obscures

propoduem; arolia present; metasomal lateral spines absent, except male T7

trifid, median spine larger than lateral spine; gonostylus of male expanded

apically, slightly notched on outer margin; penis valves reduced apically,

widely separated (Figure 15b).

Genus Pleisianthidium Cameron

Subgenus Carinanthidium Pasteels

Carinthanthidium Pasteels, 1969: 42-43. Type species: Megachile

cariniventris Friese, 1904, by original designation.

Plesianthidium (Carinthanthidium) Pasteels; Michener, 2000: 508.

Subgenus Plesianthidium Cameron s. str

Pleisianthidium Cameron, 1905: 256-257. Type species: Pleisianthidium

fulvopilosum Cameron, 1905, by original designation.

Pleisianthidium (Pleisianthidium) Cameron: Michener and Griswold, 1994:

301, 319; Michener, 2000: 508.

Subgenus Spinanthidiellum Pasteels

Spinanthidium (Spinanthidiellum) Pasteels, 1969: 55, 59-60. Type species:

Anthidium volkmanni Friese, 1909, by original designation; Pasteels,

1984: 128-129.

118

Pleisanthidium (Spinanthidiellum) Pasteels: Michener, 2000: 508.

Subgenus Spinanthidium Mavromoustakis

Dianthidium (Spinanthidium) Mavromoustakis, 1951: 977-980. Type species:

Anthidium trachusiforme Friese, 1913, by original designation.

Pleisianthidium (Spinanthidium) Mavromoustakis: Michener, 2000: 508-509.

Comment

This genus was first described by Cameron (1905). Since then, other

authors, such as Mavromoustakis (1951), Pasteels (1969, 1984) and Michener

and Griswold (1994), have revised the group. Michener (2000) listed four

subgenera, all of which are endemic to South Africa.

Diagnosis


5.4-8.2 mm; body 7.3-12.9 mm. Vestiture: white (in Spinanthidium), yellow or

reddish-brown (in nominative subgenus and Spianthidiellum), black (in

Carinanthidium); moderate to dense; female scopa on S2-S5; male S5 with

sternal comb, except in Spinanthidium. Integument: black, except brown

integumental bands in Spinanthidium (Plate 17a); female mandible with four

dissmilar teeth; lower clypeal margin without projection; subantennal suture

straight; juxtantennal carina absent; preoccipital ridge carinate dorsally; scuto-

scutellar suture closed; similar to scuto-axilla suture; scutellum usually

rounded posteriorly; arolia present; metasomal lateral spines absent; T7 of

male trifid (Plate 18c); gonostylus expanded apically; penis valves narrowly

separated (Figure 15c; Plate 18a).

119

Plate 17a and b. Two genera of the tribe Athidiini. (a) Pleisianthidium

fulvopilosum (Cameron). (b) Rhodanthidium siculum (Spinola).

Plate 18a, b and c. Some feactures used for the diagnosis of the genera. (a)

Male genitalia of P. fulvopilosum (Cameron). (b) Male genitalia of R. siculum

(Spinola). (c) Arrows indicating the trifid nature of tergite 7 of the male of R.

siculum (Spinola).

a

b

a b

c

120

Genus Pseudoanthidium Smith

Subgenus Exanthidium Pasteels

Exanthidium Pasteels, 1969: 82. Type species: Anthidium eximum Giraud,


Pseudoanthidium (Exanthidium) Pasteels: Michener and Griswold, 1994: 320;


Subgenus Micranthidium Cockerell

Pachyanthidium (Micranthidium) Cockerell, 1930b: 45-46, 48. Type species:

Anthidium truncatum Smith, 1854, by original designation.

Pseudoanthidium (Micranthidium) Cockerell: Michener, 2000: 510.

Subgenus Pseudoanthidium Friese s. str.

Anthidium (Pseudoanthidium) Friese, 1898: 101. Type species: Anthidium

alpinum Morawitz, 1873, designated by Sandhouse, 1943: 593.

Pseudoanthidium (Pseudoanthidium) Pasteels: Michener and Griswold, 1994;


Paranthidiellum Michener, 1948: 25. Type species; Anthidium cribratum

Morawitz, 1875, by original designation; Michener and Griswold, 1994:

320, syn.

Pseudoanthidium (Paraanthidiellum) [!] Michener: Pasteels, 1969: 79.

Psuedoanthidium (Carinellum) Pasteels, 1969: 80. Type species: Anthidium

ochrognathum Alfken, 1932, by original designation; Pasteels, 1969: 80,

syn.

Trachusa (Orientotrachusa) Gupta, 1993: 50. Type species: Anthidium

orientale Bingham, 1897, by original designation; Michener, 2000: 510,

syn.

121

Subgenus Royanthidium Pasteels

Royanthidium Pasteels, 1969: 86. Type species: Anthidium melanurum Klug,


Reanthidium Pasteels, 1969: 87. Type species: Anthidium reticulatum

Mocsáry, 1884, by original designation; Michener and Griswold, 1994:

320, syn.

Pseudoanthidium (Royanthidium) Pasteels: Michener and Griswold, 1994:

320; Michener, 2000: 510.

Subgenus: Semicarinella Pasteels

Pseudoanthidium (Semicarinella) Pasteels, 1984: 32. Type species:

Pseudoanthidium lattarse Pasteels, by original designation; Michener and


Subgenus Tuberanthidium Pasteels

Tuberanthidium Pasteels, 1969: 87-88. Type species: Anthidium

tuberculiferum Brauns, 1905, by original designation; Pasteels, 1984:

60-63; Gess, 1996: 290.

Pseudoanthidium (Tuberanthidium) Pasteels: Michener and Griswold, 1994:

320; Michener, 2000: 510-511.

Comment

Genus Pseudoanthidium was first described by Friese (1898). Michener

(2000) recorded six subgenera, two of which are endemic to Africa, three are

known from the Palaearctic and Orient regions. The other subgenus occurs in

both Africa and Arabia.

122

Diagnosis


4.2-7.1 mm; body 5.2-9.4 mm. Vestiture: white to reddish-black; sparse to

impubescent; female scopa on S2-S6; male sternal comb on S3. Integument:

black, with extensive yellow or reddish maculations on most of lower half of

face, lateral regions of T1-T5 (yellow areas increase progressively from front

to back), T6; Tuberanthidium mostly black; mandible of female with 5 or

more teeth; lower clypeal margin without projection; subantennal suture

arcuate; juxtantennal carina absent; preoccipital area carinate laterally; scuto-

scutellar suture slightly open, forming shiny fovae; scuto-axilla suture closed;

scutellum, in dorsal view, truncate posteriorly; arolia absent; metasomal lateral

spines absent; gonostylus of male slightly expanded apically; penis valve

narrow apically (Figure 16a).

123

Rhodanthidium

Figure 16a, b, c and d. Male genitalia of four anthidine bees. (a)

Pseudoanthidium truncatum (Smith). (b) Rhodanthidium siculum (Spinola).

(c) Serapista denticulata Cockerell. (d) Trachusa aquiphilum (Strand).

b a

c

d

124

Genus Rhodanthidium Spinola

Subgenus Asianthidium Popov

Meganthidium (Asianthidium) Popov, 1950: 315. Type species: Anthidium

glasunovii Morawitz, 1894, by original designation.

Rhodanthidium (Asianthidium) Popov: Michener and Griswold, 1994: 322;


Rhodanthidium (Trianthidium) Mavromoustakis, 1958: 435. Type species:

Anthidium caturigense Giraud, 1863, by original designation;

Warncke, 1980: 119-209, syn.

Meganthidium (Oxyanthidium) Mavromoustakis, 1963a: 653. Type species:

Anthidium aculeatum Klug, 1832, by original designation; Warncke,

1980: 119-209, syn.

Axillanthidium Pasteels, 1969: 39. Type species: Axillanthidium axillare

Pasteels, 1969, by original designation; Warncke, 1980: 119-209, syn.

Subgenus Meganthidium Popov

Meganthidium Popov, 1950: 315. Type species: Anthidium christophi

Morawitz, 1876, by original designation; Pasteels, 1969: 31.

Rhodanthidium (Meganthidium) Popov: Michener and Griswold, 1994: 322;

Michener, 2000: 511-512.

Subgenus Rhodanthidium Isensee s. str.

Anthidium (Rhodanthidium) Isensee, 1927; 374. Type species: Anthidium

siculum Spinola, 1838, by original designation.

Bellanthidium Pasteels, 1969: 38. Type species: Anthidium infuscatum

Erichson, 1835, by original designation; Michener and Griswold, 1994:

322, syn.

125

Rhodanthidium (Rhodanthidium) Isensee: Michener and Griswold, 1994: 322;


Comment

The genus Rhodanthidium was first described by Iseense (1927).

Michener (2000) recognized three subgenera, all of which occur in the

Palaearctic Region, however, one also occurs in the Oriental Region.

Diagnosis


9.2-11.2 mm; body 11.2-17.1 mm. Vestiture: white to reddish-brown, sparse to

moderate, very dense on legs; female scopa on S2-S5; male S4-S5 with medial

sternal comb. Integument: mostly black, (Meganthidium mostly yellow) with

extensive yellow to reddish brown maculations on face and metasoma (Pl.

12b); mandible of female with four dissimilar, large teeth; lower clypeal

margin without projection; subantennal suture more or less straight;

juxtantennal carina absent; preoccipital area gently rounded; scuto-scutellar

suture closed, similar to scuto-axilla suture; scutellum, in dorsal view, rounded

posteriorly; arolia present; metesoma with reduced lateral spines; T7 of male

trifid (Pl. 12e); male gonostylus slender with outer margin deeply notched;

penis vales narrowly seperated, with their apices curved inward (Figure 16b;

Pl. 12d).

126

Genus Serapista Cockerell

Serapis Smith, 1854: 218-219 (not Link, 1839). Type species: Serapis

denticulatus Smith, 1854, monobasic.

Serapista Cockerell 1904b: 357. Replacement for Serapis Smith, 1854;

Cockerell 1932: 360; Cockerell 1933c: 357; Cockerell 1935: 237;

Mavromoustakis, 1940a: 65; Pasteels 1969: 107; Pasteels 1984: 38;


Comment

Serapista--This robust, megachiliform to chalicodomiform, pollen-

collecting genus was first described by Smith (1854) as Serapis. However,

because Serapis was preoccupied under phyllum mollusca, Cockerell (1904)

renamed this genus as Serapista Cockerell. Other authors, such as

Mavromoustakis (1940) and Pasteels (1969, 1984), worked on this genus.

Pasteels (1984) recorded four species all of which are endemic to Africa.

Diagnosis


6.7-8.1 mm; body 9.8-11.5 mm. Vestiture: mostly white with unique lateral

bands on T1-T6 (increasing in size pregressively from T1 to T5, reduced on

T6, Plate 19a); brownish on legs; sparse to impubescent; female scopa on S2-

S5; male sternal comb absent. Integument: entirely black, female mandible

with eleven dissimilar teeth; lower clypeal margin with six projections;

subantennal suture straight; juxtantennal carina well developed;

mediolongitudinal region of face

127

strongly crinate; preoccipital area gently rounded; scuto-scutellar suture

closed, similar to scuto-axilla suture; posterior region of axilla with strong

spine (Plate 20a); scutellum, in dorsal view, truncate posteriorly; arolia absent;

female T1-T6, male T1-T6 with lateral spines (Plate 19a); T7 of male trifid

(Plate 20b); gonostylus narrows anteriorly; penis valve arcuate in profile,

notched anteriorly, widely seperated (Figure 16c).

128

Plate 19a and b. Two anthidine bees. (a) Arrow indicating the lateral spines on

the metasoma of Serapista denticulata Cockerell. (b) Female of Trachusa

aquiphilum (Strand).

Plate 20a and b. Two features used for diagnosis of Serapista denticulata

Cockerell. (a) Arrows indicating lateral extension of axilla. (b) Arrows

showing the trifid nature of tergite 7 of the male.

a

b

d c

129

Genus Stelis Panzer

Subgenus Dolichostelis Parker and Bohart

Dolichostelis Parker and Bohart, 1979: 138. Type species: Stelis laticincta,

Cresson, 1878, by original designation.

Stelis (Dolichostelis) Parker and Bohart: Michener, 2000: 513.

Subgenus Malanthidium Pasteels

Malanthidium Pasteel, 1969: 26. Type species: Anthidium malaccense Friese,


Stelis (Malanthidium) Pasteels: Michener and Griswold, 1994: 322-323;


Subgenus Protostelis Friese

Protostelis Friese, 1895: 25. Type species: Stelis: freygessneri Friese, 1885, by

designation of Popov, 1938.

Stelis (Protostelis) Friese: Michener, 1944: 258-259, syn; Michener, 1948: 4;


Heterostelis Timberlake, 1941: 125. Type species: Stelis anthidiodes

Timberlake, 1941, by original designation; Griswold and Michener,

1988: 37, syn.

Doxanthidium Pasteels, 1969: 28. Type species: Anthidium paradoxum

Mocsáry, 1884, by original designation; Pasteels, 1969: 28, syn

Subgenus Pseudostelis Popov

Pseudostelis Popov, 1956: 167. Type species: Stelis strandi Popov, 1935, by

original designation.

Stelis (Pseudostelis) Popov: Michener and Griswold, 1994: 322; Michener,

2000: 515.

130

Subgenus Stelidomorpha Morawitz

Stelidomorpha Morawitz, 1875: 131. Type species: Anthidium nasutum

Latreille, 1809, monobasic.

Stelis (Stelidomorpha) Morawitz: Michener and Griswold, 1994: 322;


Subgenus Stelis Panzer s. str.

Trachusa Jurine, 1801: 164 (not Panzer 1801). Type species: Apis aterrima

Panzer, 1798, designated by Morice and Durrant, 1915: 426.

Suppressed by commission Opinion 135, 1939 (Direction 4).

Stelis Panzer, 1806: 264. Type species: Apis aterrima Panzer, 1798 (not

Christ, 1791), monobasic. Isogenotypic with Trachusa Jurine.

Stelis (Stelis) Panzer: Michener and Griswold, 1994: 322; Michener, 2000:

515-516.

Gyrodroma Klug, 1807: 198. Type species: Apis aterrima Panzer, 1798

[Sandhouse incorrectly considered Gyrodroma to be monobasic,

however, two species were listed by Klug, 1807: 225]. Isogenotypic

with Stelis.

Gymnus Spinola, 1808: 9. Type species: Apis aterrima Panzer, 1798.

Isogenotypic with Stelis.

Ceraplastes Gistel, 1848: x, unjustified replacement name for Stelis Panzer,

1806. Type species: Apis aterrima Panzer, 1798. Isogenotypic with

Stelis.

Chelynia Provancher, 1888: 296, 322. Type species: Chelynia labiata

Porvancher, 1888, monobasic; Michener, 1944: 258, syn; Michener,

131

1997: 15.

Melanostelis Ashmead, 1898: 283. Type species: Melanostelis betheli

Ashmead, 1898, by original designation; Michener, 1948: 4, syn.

Stelidium Robertson, 1902: 323. Type species: Stelidium trypetinum

Robertson, 1902, monobasic; Robertson, 1902: 323, syn.

Microstelis Robertson, 1903: 170, 175. Type species: Stelis lateralis Cresson,

1864, by original designation; Michener, 1944, syn; Michener 1948:

4; Michener 2000: 515.

Stelis (Pavostelis) Sladen, 1916: 313. Type species: Stelis montana Cresson,

1864, monobasic; Michener, 1944: 258, syn; Michener, 1948: 4.

Stelis (Stelidina) Timberlake, 1941: 131. Type species: Stelis hemirhoda

Linsley, 1939, by original designation; Michener, 1944: 258, syn;

Michener, 1948:4.

Stelis (Stelidiella) [!] Timberlake: Michener, 1944: 258-259.

Stelis (Stelidium) Robertson, 1902: 323. Type species: Stelidium trypetinum

Robertson, 1902, by original designation; Michener, 1944: 258-259,

syn; Michener, 1948: 4.

Stelis (Leucostelis) Noskiewicz, 1961: 126, 132. Type species: Gyrodroma

ornatula Klug, 1807, by original designation; Noskiewicz: 1961:

132, syn.

Comment

This cleptoparasitic genus was first described by Panzer (1806). Since

then several other authors, such as Provancher (1888), Ashmead (1898),

132







record a Kenyan species but did record one species from South Africa. This

genus was not available for examination during this study, due to the fact that,

it is a rare specimen and the museum was unwilling to loan them out.

Genus Trachusa Panzer

Subgenus Archianthidium Mavromoustakis

Archianthidium Mavromoustakis, 1939: 91. Type species: Anthidium laticeps

Morawitz, 1873, by original designation.

Trachusa (Archianthidium) Mavromoustakis: Michener and Griswold, 1994:

323; Michener, 2000: 517.

Subgenus Congotrachusa Pasteels

Trachusa (Congotrachusa) Pasteels, 1969: 24. Type species: Anthidium

schoutedeni Vachal, 1901, by original designation; Pasteels, 1984:

117; Michener and Griswold, 1994: 323; Michener, 2000: 517.

Subgenus Hetereanthidium Cockerell

Anthidium (Heteranthidium) Cockerell, 1904b: 292. Type species: Anthidium

dorsale Lepeletier, 1841, by original designation.

Trachusa (Heteranthdium) Cockerell: Griswold and Michener, 1988: 27-28;

133

Michener, 2000: 517-518.

Subgenus Legnanthidium Griswold and Michener

Trachusa (Legnanthidium) Griswold and Michener, 1988: 27. Type species:

Anthidium ridingsii Cresson, 1878, by original designation; Michener,

2000: 518.

Subgenus Massanthidium Pasteels

Trachusa (Massanthiidium) Pasteels, 1969: 24. Type species: Trachusa

flavorufula Pasteels, 1969, by original designation; Pasteels, 1984:

118; Michener and Griswold, 1994: 323; Michener, 2000: .518.

Subgenus Metatrachusa Pasteels

Trachusa (Metatrachusa) Pasteels, 1969: 22. Type species: Anthidium

pendleburyi Cockerell, 1927, by original designation; Michener and


Subgenus Orthanthidium Mavromoustakis

Paraanthidium (Orthanthidium) Mavromoustakis, 1953: 837. Type species:

Anthidium formosanum Friese, 1917, by original designation.

Trachusa (Orthanthidium) Mavromoustakis: Pasteels, 1969: 23; Michener and


Subgenus Paraanthidium Friese

Anthidium (Paraanthidium) Friese, 1898: 101. Type species: Apis interrupta

Fabricius, 1781, designated by Cockerell, 1909: 269.

Trachusa (Paraanthidium) Friese: Pasteels, 1984: 114-115; Michener and

134

Griswold, 1994: 323; Michener, 2000: 518-519.

Protanthidium Cockerell and Cockerell, 1901: 49. Type species: Megachile

steloides Bingham, 1896, by original designation; Mavromoustakis,

1939: 90, syn.; Michener, 1948: 23, 26.

Trachusa (Philotrachusa) Pasteels, 1969: 22. Type species: Anthidium

aquilifilum Strand, 1912, by original designation; Pasteels: 1969: 22,

syn.

Subgenus Trachusa Panzers. Str.

Trachusa Panzer, 1804: 14-15. Type species: Trachusa serratulae Panzer,

1804, designated by Sandhouse, 1943: 605.

Trachusa (Trachusa) Panzer: Pasteels, 1969: 21; Michener and Griswold,

1994: 325; Michener, 2000: 519.

Diphysis Lepeletier, 1841: 307. Type species: Diphysis pyrenaica Lepeletier,

1841 (= Trachusa serratulae). Isogenotypic with Trachusa.

Megachileoides Radoszkowski, 1874: 132. Type species: Trachusa serratulae

Panzer, 1804, designated by Michener, 1995: 375; Isogenotypic with

Trachusa.

Megachiloides [!] Radoszkowski: Saussure, 1890: 35; Michener, 2000: 519.

Subgenus Trachusomimus Popov

Trachusomimus Popov, 1964: 406. Type species: Trachusa perdita Cockerell,


Trachusa (Trachusomimus) Popov: Griswold and Michener, 1988: 25;


135

Subgenus Ulanthidium Michener

Ulanthidium Michener, 1948: 13. Type species: Ulanthidium mitchelli

Michener, 1948, by original designation.

Ulanthidium (Olmecanthidium) Peters, 1972b: 377. Type species:

Ulanthidium interdisciplinaris Peters, 1972, by original designation;

Peter, 1972: 13, syn.

Trachusa (Ulanthidium) Michener: Griswold and Michener, 1988: 26;


Comment

This non-parasitic genus was first described by Panzer in 1806. Since

then several other authors, such as Provancher (1888), Ashmead (1898),








Diagnosis


6.1-8.4 mm; body 9.0-11.6 mm. Vestiture: pale white to brownish; sparse to

moderately dense pubescent; female scopa on S2-S5; male sternal comb on

S4-S5. Integument: mostly black, with limited to extensive pallid maculation

136

on face, scutellum and metasoma (Plate 19b); female mandible with four

dissimilar teeth, sometimes five or six, as in subgenera Massanthidium and

Congotrachusa; lower clypeal margin with six projections; subantennal suture

straight; juxtantennal carina absent; preoccipital area gently rounded; scuto-

scutellar suture closed, similar to scuto-axilla suture; scutellum, in dorsal

view, truncate or emarginate postero-medially; arolia absent, sometimes

reduced in subgenus Congotrachusa; metasomal without lateral spine;

gonostlyus expanded apically, deeply notched on inner margin; penis valves

reduced in size and narrowly separated (Figure 16d).

Genus Trachusoides Michener and Griswold

Trachusoides Michener and Griswold, 1994: 324. Type species: Trachusoidse

simplex Michener and Griswold, 1994, by original designation;


Comment

This non-parasitic, large megachiliform genus was first described by

Michener and Griswold (1994). This monotypic genus is endemic to southern

India. The representative species of this genus was not available for

examination during this study, due to the fact that, it is a rare specimen and the

museum was unwilling to loan it out.

137

CLADOGRAMS AND TREE TOPOLOGIES OF GENUS Anthidiellum

In the cladistic analysis of the genus Anthidiellum with heuristic

search, 385 most parsimonious trees were found, with L: 39 steps, CI: 79%

and RI: 81%. Figure 17 depicts a typical cladogram topology of most of the

385 most parsimonious trees. The strict consensus tree, on the other hand, had

L: 51, CI: 60%, RI: 53% collapsed 14 unsupported nodes (Fig. 18). The strict

consensus values using bootstrap and jackknife search tool were found to be:

L: 48/ 48, CI: 64/ 64%, RI: 60/ 60% (Figure 19-20).

The analyses of the ground-plan of those trees revealed three major

clades outside the outgroups (Trachusa and Eoanthidium) and these consist of:

(1) Anthidiellum flavescens and A. rubellum. 2) A. eritrinum, A. absonulum, A.

apicatum, A. auriscopatum, A. frontoreticulatum, A. kimberleyana, A.

orichalciscopatum, A. polyochrum, A. krugerense, A. sternale, A.

tegwaniense, A. zebra, A. bulawayense, A. otavicum, A. bipectinatum, A.

spilotum. The probable phylogenetic implications have been dealt with in the

discussion.

138

Trachusa aquiliphilum

Eoanthidium turnericum

Anthidiellum absonulum

A. apicatum

A. auriscopatum

A. bipectinatum

A. bulawayense

A. eritrinum

A. rubellum

A. flavescens

A. frontoreticulum

A. kimberleyana

A. orichalciscopatum

A. otavicum

A. polyochrum

A. Pygn. sp1

A. spilotum

A. sternale

A. tegwaniense

A. zebra

29

1

16

1

7

1

5

1

4

1

0

-

0

-0

-

13

1

0

-

0

-

5

0

0

-

0

-

0

-

21

1

18

0

17

1

16

1

12

1

8

2

0

-

0

-

0

-

0

-

0

-

0

-

20

1

0

-

0

-

0

-

0

-

17

1

0

-

0

0

26

0

0

0

0

22

2

0

0

13

1

0

0

0

5

1

0

26

1

0

0

9

1

2

1

0

0

26

1

15

1

8

1

1

1

0

27

1

22

1

14

1

11

1

10

1

6

1

0

0

Figure17. Cladogram indicating phylogeny of the Afrotropical species of

genus Anthidiellum of the heuristic analysis.

139

Figure 18. Cladogram indicating the strict concensus tree of the Afrotropical

species of genus Anthidiellum



A. absonulum

A. apicatum

A. auriscopatum

A. bipectinatum

A. bulawayense

A. eritrinum

A. rubellum

Anthidiellum flavescens

A. frontoreticulum

A. kimberleyana


A. otavicum

A. polyochrum

A. krugerense

A. spilotum

A. sternale

A. tegwaniense

A. zebra

83

70

70

92

90

99

100

140

Figure 19. Cladogram indicating phylogeny of the Afrotropical species of

genus Anthidiellum from the bootstrap analysis



A. absonulum

A. apicatum

A. auriscopatum

A. bipectinatum

A. bulawayense

A. eritrinum

A. rubellum


A. frontoreticulatum

A. kimberleyana


A. otavicum

A. polyochrum

A. krugerense

A. spilotum

A. sternale

A. tegwaniense

A. zebra

73

68

70

94

92

99

100

141

Figure 20. Cladogram indicating phylogeny of the Afrotropical species of

genus Anthidiellum from the jackknife analysis



A. absonulum

A. apicatum

A. auriscopatum

A. bipectinatum

A. bulawayense

A. eritrinum

A. rubellum


A. frontoreticulatum

A. kimberleyana


A. otavicum

A. polyochrum

A. krugerense

A. spilotum

A. sternale

A. tegwaniense

A. zebra

73

70

70

95

89

100

100

142

SYSTEMATIC REVISION OF THE AFROTROPICAL SPECIES OF

THE GENUS Anthidiellum (COCKERELL)


Anthidium (Anthidiellum) Cockerell, 1904c: 3. Type species: Trachusa


Anthidiellum Cockerell: Pasteels, 1969: 45; Pasteels, 1984: 92; Michener and


Comment

This genus was described as a subgenus of Anthidium and later given

generic status by Pasteels (1969). However, the taxon appears be defined by

only two homoplasious apomorphic characters within the tribe Anthidiini. Its

arcuate subantennal suture and fovea of the scuto-axilla suture are also found

in Bathanthidium and Eoanthidium, though fovea is narrow in these genera, as

well as in Acanthidium. The fovea that is margined by a carina behind the

propodeal spiracle differs from that in Duckeanthidium and Stelis. However,

results in the analysis reveals two major clades of trees within the genus. This

apparently is not compatible with the subgeneric classification of genus by

Michener (2000). For the purpose of this study, the subgeneric classification

of Michener (2000) has been retained.

The Afrotropical genus comprises three subgenera. Two of these

subgenera are endemic species, while the third subgenus occurs in both Orient

and Afrotropical regions. 17 species are currently known to occur in

Afrotropical Region.

143

Keys to the Species

A taxonomic key is provided below for all the Afrotropical species that were

available for examination during this study.

1. Mandible with three teeth, apical region narrow; antennal scape

long, subequal 7X first flagella segment (male) -----------------------------2.

— Mandible with four teeth, apical region broad; antennal scape short, 3X

first flagella segment (female) -------------------------------------------------15.

2. Tegula broad anteriorly, lateral margin convex; pronotal lobe lamella------3.

— Tegula narrows anteriorly, lateral margin concave; pronotal lobe carinate

mesally, becoming progressively lamellate distally---------------------------5.

3. Sternal comb present medially on S5, head, mesosoma black, metasoma

yellowish brown --------------------------------------.Anthidiellum flavescens.

— Sternal comb absent, integument mostly reddish brown to black with

yellow integumental maculations -----------------------------------------------4.

4. Mesosoma entirely black, with large, very dense punctures; 12 hamuli on

hind wing--------------------------------------------------------------A. eritrinum.

— Mesosoma black, except yellow to reddish brown integumental maculation

on axilla and scutellum, punctations small and moderately dense; 15

hamuli on hind wing -------------------------------------------------A. rubellum.

5. Gonostylus narrows apically------------------------------------------------------6.

— Gonostylus broadens apically ----------------------------------------------------7.

6. Penis valve broad apically ----------------------------------------A. tegwaniense.

—. Penis valves narrow apically -------------------------------------------.A. zebra.

7. Ventral margin of clypeus with four projections -------------------------------8.

144

— Ventral margin of clypeus with five to six projections ----------------------11.

8. Hind wing with 10 hamuli on anterior margin ---------------------A. otavicum.

— Hind wing with 11 or more hamuli on anterior margin -----------------------9

9. Sternal comb very long (about 40 setae long) ----------------A.

frontoreticulatum.

— Sternal comb short (12 or less) -------------------------------------------------10.

10. Penis valve narrow apically, gonostylus slightly notched on outer margin -

---------------------------------------------------------------------------A. spilolum.

— Penis valve broad apically, gonostylus deeply notched on outer margin------

---------------------------------------------------------------------------.A. sternale.

11. Hind wing with 10 hamuli on anterior margin --------------A. bulaweyense.

— Hind wing with 11 or more hamuli on anterior margin ---------------------12.

12. Hind basitarsus slender, subequal half width of hind tibia ---A. absonulum.

—Hind basitarsus as broad as hind tibia-------------------------------------------13.

13. Vein cu-v of hind wing oblique (slants) -----------------------A. polyochrum.

— Vein cu-v of hind wing perpendicular (straight) -------------A. bipectinatum.

14. Tegula broad anteriorly, lateral margin convex; pronotal lobe lamellate;

distance between antennal sockets narrow, subequal two antennal socket

diameters -------------------------------------------------------------------------15.

— Tegula narrows anteriorly, lateral margin concave; pronotal lobe carinate

mesally, becoming progressively lamellate distally; distance between

antennal sockets wide, more than two antennal socket diameters --------17.

15. Preoccipita1 area carinate laterally; mesosoma yellowish brown; scutellum

145

(in dorsal view) truncate posteriorly -----------------------------A. flavescens.

— Preoccipital area carinate dorso-laterally; mesosoma reddish brown to

black with yellow or reddish integumental maculations; scutellum (in

dorsal view), emarginate medio-posteriorly --------------------------------16.

16. Mesosoma entirely black, with large, very dense punctures; 12 hamuli on

hind wing ------------------------------------------------------------.A. eritrinum.

— Mesosoma black, except yellow to reddish brown integumental maculation

on axilla and scutellum, punctations small and moderately dense; 15

hamuli on hind wing -------------------------------------------------A. rubellum.

17. Ventral margin of clypeus with four projections----------------------------18.

— Ventral margin of clypeus with five or more projections ------------------27.

18. Mandible with outer carina only -----------------------------------------------19.

— Mandible with both outer and inner carinae ----------------------------------22.

19. Hind wing with 10 hamuli on anterior region --------------------A. otavicum.

— Hind wing with 11 or more hamuli --------------------------------------------20.

20. Vein cu-v of hind wing oblique (curves back); 11 hamuli on hind wing ----

-----------------------------------------------------------------A. frontoreticulatum.

— Vein cu-v of hind wing perpendicular (straight); 12 hamuli on hind wing---

--------------------------------------------------------------------------------------21.

21. Hind basitarsus slender, subequal half as broad as hind tibia -----------------

----------------------------------------------------------------------A. tegwaniense.

— Hind basitarsus broad, as wide as hind tibia or broader than hind tibia.------

--------------------------------------------------------------------A. auriscopatum.

146

22. Hind wing with eleven hamuli on anterior margin -------------------------23.

— Hind wing with twelve hamuli on anterior region -----A. orichalciscopatum.

23. Vein cu-v of hind wing straight.------------------------------------------------24.

— Vein cu-v of hind wing oblique (curves back) -------------------------------25.

24. Tibiae with thick stout setae.---------------------------------------------A. zebra.

— Tibiae lack thick stout setae ----------------------------------------.A. apicatum.

25. Scutum (in dorsal view) extend laterally and longitudinally, forming

groove with carina next to tegula----------------------------------A. krugerense

— Scutum (in dorsal view) rounded, lack lateral extension --------------------26.

26. Tibiae with thick stout setae; mandible with very sharp outer teeth ----------

----------------------------------------------------------------------------A. sternale.

— Tibiae lacks thick stout setae; mandible with blunt outer teeth ----------------

---------------------------------------------------------------------------A. spilotum.

27. Mandible with outer carina only.------------------------------A. bulawayense.

— Mandible with both outer and inner carinae ----------------------------------28.

28. Vein cu-v of hind wing straight ------------------------------------------------29.

— Vein cu-v of hind wing oblique (curves back) -------------------------------30.

29. Hind wing with 11 hamuli.-----------------------------------------------A. zebra

— Hind wing with 12 hamuli--------------------------------------.A. kimberleyana.

30. Mandible with entirely large punctures ------------------------A. polyochrum.

— Mandible with large punctures basally, small punctures apically -------------

----------------------------------------------------------------------A. bipectinatum.

147

Subgenus Anthidiellum Cockerell s. str.

Anthidium (Anthidiellum) Cockerell, 1904c: 3. Type species: Trachusa


Anthidium (Cerianthidium) Friese, 1932: 304. Type species: Trachusa strigata

Panzer, 1805. Isogenotypic with subgenus Anthidiellum.

Anthidiellum (Anthidiellum) Cockerell: Pasteels, 1969: 45; Pasteels, 1984: 92;


Comment

This subgenus comprises two Afrotropical species, Anthidiellum

eritrinum (Friese) and Anthidiellum rubellum (Friese). A. eritrinum is endemic

Eritrea, whereas A. rubellum is only known from South Africa.

General Diagnosis of the Genus


3.8-5.4 mm; body 5.4-5.8 mm. Vestiture: cream to reddish brown; sparse to

impubescent; male sternal comb absent; upper edge of inner surface of hind

tibia not evidently marked by limited area of keirotrichiate. Integument: black,

except yellow or reddish maculation on most of face, entire margin of

preoccipital area, margins of scutum, axilla and scutellum, lateral regions of

T1-T4 (in A. eritrinum); preoccipital carina present dorso-laterally; paraocular

carina well developed; vertex convex, 2X ocellus diameters above eye;

distance between antennal sockets subequal two antennal sockets diameter;

subantennal suture strongly curved, subequal clypeus length; scape long, 5X

148

length of first flagellar segment; antennal flagellum of male long (0.70-0.84

head length); apex of female mandible twice as broad as base; pronotal lamella

well developed, extends mesad well past edge of scutum; tegula not narrowed

anteriorly, lateral margin convex; omaular distinct, weakly lamellate,

extending ventrally, not reaching midline; axilla extending laterally beyond

lateral margin of scutum; rows of pits across upper margin of propodeum

weakly developed, not carinate; corbiculate sculpture on propodeum confined

to basolateral region; entire surface of fore wing with numerous hair and

papillae; jugal lobe of hind wing about half as long as vannal lobe; T2-T5, and

T6 in dorsolateral view, gently curved towards apex; T5-T6 in female, T6-T7

of males curved weakly downward and appears swollen.

Anthidiellum (Anthidiellum) eritrinum (Friese)

Anthidium eritrinum Friese, 1915: 275, 295-296, female holotype (ZMHB)

Eritrea; Friese, 1915: 275, 295-296.

Anthidiellum eritrinum (Friese): Mavromoustakis, 1936a: 44, 45.

Anthidiellum (Anthidiellum) eritrinum (Friese): Pasteels, 1984: 92-93, 95, 156.

Diagnosis

Male. Unknown. Female. Length: head 2.1 mm; scutum and scutellum

2.3 mm; forewing 5.4 mm; body 5.8 mm. Vestiture: white to pale yellow,

except orange on vertex, mesosomal dorsum and under tarsi; mostly sparse,

tibiae with fairly dense pubescent, without thick, stout setae. Integument: head

black with postero-lateral regions of vertex and lower face (Plate 22a) mostly

149

yellow; mesosoma mostly black, pronotal lobe and tegula partly yellow, legs

almost entirely yellow (blackish proximally and orangish distally); metasoma

black to reddish-black with following areas yellow: dorso-lateral region of T1,

wide cross-bands on T2-T5, most of T6 (Plate 21a; Figure 21a); lower clypeal

margin with five projections (Figure 21a); mandible with large punctures

basally (shiny between punctures), fine sculpture near apex, two ventro-distal

sutures; two dissimilar, large outer teeth, two very small inner teeth; lateral

margin of scutum, dorsal view, straight; scuto-scutellar suture open, forming

narrow shiny fovea; scutellum, dorsal view, generally gently convex

posteriorly, concave postero-medially, obscures propodeum; 12 long hamuli

on hind wing; vein cu-v slants; hind basitarsus narrow, more than half as wide

as hind tibia; T2, dorsal view, gently convex laterally.

Distribution

Anthidiellum eritrinum (Friese) is endemic to Eritrea.

150

Plate 21a and b. Two species of genus Anthidiellum. (a) Female of

Anthidiellum eritrinum Friese. (b) Female of Anthidiellum flavescens (Friese).

Plate 22a and b. Photographs of the faces of two species of genus

Anthidiellum. (a) Arrows indicating areas of yellow maculation on the face of

A. eritrinum Friese. (b) Arrow indicating area of yellow maculation on the

face of Anthidiellum flavescens (Friese).

a b

a b

151

Figure 21a and b. Two features used for diagnosis of Anthidiellum eritrinum

Friese. (a) Face and (b) Abdomen with arrow indicating regions of yellow

maculations

A. rubellum

Figure 22a, b, c and d. Diagnostic features of Anthidiellum rubellum Friese. (a

and b) Face, with arrow indicating regions of yellow maculations of the

female and male respectively. (c) Mesosoma (in dorsal view), arrows showing

areas of yellow maculation and lateral extension of axilla beyond scutum

margin. (d) Male genitalia (arrow showing distinctly curved nature of the

gonostylus).

a b

c d

a b

152

Anthidiellum (Anthidiellum) rubellum (Friese)

Anthidium rubellum Friese, 1905e: 74, female holotype (TMSA) South Africa;

Friese, 1909a: 400, 401, 412; Anonymous, 1958: 33.

Anthidiellum (Anthidiellum) rubellum (Friese): Pasteels, 1969: 45.

Diagnosis

Female. .Length: head 2.2 mm; scutum and scutellum 1.9 mm;

forewing 4.0 mm; body 5.9 mm. Vestiture: white, except reddish to black on

fore tarsi; sparse to impubescent; thick stout setae absent on tibiae.

Integument: head, mesosoma black, metasoma brownish, except yellow or

reddish-brown maculation on clypeus (ventral margin black, Figure 22a),

ventral region of supraclypeus, entire paraoccular area, preoccipital region and

gena, dorso-lateral surface of scutum, entire axilla, most of scutellum (antero-

median region black), dorsal region of mesepisternum and mesepimeron,

entire legs; mandible with large punctures basally, small punctures apically,

shiny between punctures, with outer and inner carinae, two, similar, large

outer teeth and two, similarly very reduced teeth; clypeus margin with three

projections on each side (one medial and one lateral); scuto-scutellar suture

open, forming shiny fovea; lateral region of sutum rounded; scutellum, in

dorsal view, truncate posteriorly, obscures propodeum (Figure 22c); 15 short

hamuli on hind wing; vein cu-v slants; hind basitarsus broad, more than half as

wide as hind tibia.

Male. Length: head 2.0 mm; scutum and scutellum 1.7 mm; forewing

3.8 mm; body 5.4 mm. Vestiture and integument: similar to female, except for

153

sex limit characters and moderately dense lateral fringes on S2-S4, S5 lacks

sternal comb; apex of mandible half as wide as in female (Figure 22b), 3 very

small teeth; gonostylus wide apically, without a notch on outer margin; penis

valves narrowly separated (Figure 22d).

Distribution

Anthidiellum rubellum Friese is endemic to South Africa.

Subgenus Chloranthidiellum Mavromoustakis

Anthidiellum (Chloranthidiellum) Mavromoustakis, 1963b: 491. Type species:

Anthidium minutulum flavescens Friese, 1925, by original designation;

Pasteels, 1984: 92; Michener and Griswold, 1994: 312; Michener,

2000: 487.

Anthidiellum (Chloranthidium) [!] Mavromoustakis: Pasteels, 1969: 48.

Comment

This subgenus is monotypic, comprising Anthidiellum flavescens

(Friese), endemic to East and southern Africa.

This study agree with the position of Anthidiellum hessi (Cockerell), as

synonym of A. flavescens based on similar morphological characters such as

integument colour and vestiture, carina on head and genitalia of male.

154

Anthidiellum flavescens (Friese)

Anthidium minutulum var. flavescens Friese, 1925a: 509, male holotype

(NMBZ) Zimbabwe.

Anthidiellum flavescens (Friese): Mavromoustakis, 1936b: 605-606; Michener,

2000: 487.

Anthidiellum (Chloranthidiellum) flavescens (Friese): Mavromoustakis,

1963b: 491; Pasteels, 1984: 93-95.

Anthidiellum (Chloranthidiellum) flavescens nigrescens Pasteels, 1984: 94,

female holotype (SEMC) Kenya.

Anthidiellum hessei Cockerell, 1932c: 523-524, male syntypes (SAMC or

NHML) Namibia; Cockerell, 1936c: 456; Mavromoustakis, 1936b:

605, syn.

Diagnosis

Female. Length: head 1.4-1.8 mm; scutum and scutellum 1.9-2.1 mm;

forewing 4.5-4.6 mm; body 5.6-5.8 mm. Vestiture: white, except reddish to

black on tarsi; sparse to impubescent; tibiae without thick, stout setae.

Integument: head and mesosoma black, except yellow maculations on most of

lower half of face (Plate 22b), a stripe on dorsal corner of preoccipital area,

spot on doral surface of axilla, entire dorsal and lateral surfaces of legs (Figure

23e); anterior metasoma orange to reddish-brown with yellow maculations on

anterior regions of T1-T6 (Figure 23b; Plate 21b); preoccipital carina present

laterally, absent dorsally; paraocular carina well developed; vertex convex, 2X

ocellus diameters above eye; distance between antennal sockets subequal two

155

antennal socket diameter; subantennal suture strongly curved, subequal to

clypeus length; scape long (first flagellar segments less than 3X scape length);

antennal flagellum short (about half head length); lower clypeal margin with

five very small papillae (one median, two lateral); apex of mandible 1.5X as

wide as base; mandible finely sculptured with small punctures, shiny between

punctures, with two carinae (outer and inner carinae), two large similar outer

teeth and two small dissimlar inner teeth; pronotal lobe strongly lamellate,

lamella tapers, extending nearly as far mesad from lateral margin of scutum as

laterad; tegula, not narrowed anteriorly, with lateral margin convex; omaular

distinct, weakly lamellate, extending ventrally, not reaching midline; axilla

extending laterally beyond lateral margin of scutum; scuto-scutellar suture

open, forming shiny fovae; scutellum, in dorsal view, truncate posteriorly,

hides median region of propodeum (Figure 23a); rows of pits across the upper

margin of propoduem strongly developed, evident more horizontally, lateral

margin with carina; 12 hamuli on hind wing; vein cu-v slants; hind barsitarsi

nearly as long as broad.

Male: Length: head 1.5-1.6 mm; scutum and scutellum 1.6-1.7 mm;

forewing 4.2-4.3 mm; body 4.8-5.0 mm. Vestiture and integument: Similar to

female (Figure 23c), except apical region of S5 with medial comb of about

thirty similar setae; clypeus entirely yellow; mandibles mostly bright yellow

with apex dark brown; gonstylus expanded apically, deeply notched on outer

margin; penis valves narrowly separated (Figure 23d).

156

Distribution

Anthidiellum flavescens (Friese) is widespread in Kenya and southern

Africa.

157

A. flavescens

A. flavescens

Figure 23a, b, c, d and e. Diagnostic features of Anthidiellum flavescens

(Friese). (a) Mesosoma (in dorsal view), arrows showing areas of yellow

maculation and lateral extension of axilla beyond scutum margin. (b and c)

Metasoma in dorsal view, with arrows indicating regions of yellow

maculations of the female and male, respectively. (d) Male genitalia. (e) Hind

leg (arrows showing regions of integumental maculations).

a

b

d

c

e

158

Subgenus Pycnanthidium Krombein

Pycnanthidium Krombein, 1951: 292. Type species: Pycnanthidium solomonis

Krombein, 1951, by original designation.

Pygnanthidium [!] Krombein, 1951: Mavromoustakis, 1963b: 491.

Pygnanthidium [!] (Pygnanthidiellum) Mavromoustakis, 1963b: 492. Type

species: Anthidium zebra, 1904, by original designation.

Anthidiellum (Pycnanthidium) Krombein: Pasteels, 1969: 45; Michener and


Anthidiellum (Pygnanthidiellum [!]) Mavromoustakis: Pasteels: 1984: 94.

Comment

This is the most diverse group among the Afrotropical subgenera.

Pasteels (1984) listed 13 Afrotropical species. This subgenus also occurs in

Asia and Australia. Currently, two Madagascan species have been included,

bringing the number of species to 15 Afrotopical species. These consist of

Anthidiellum absonulum (Cockerell)= A. aspidopodum Mavromoustakis, A.

apicatum (Smith), A. bipectinatum Pasteels, A. bulawayense Mavromoustakis,

A. madli Pauly, A. mediale Pasteels, A. micheneri Pauly, A. orichalciscopatum

(Strand) = A. auriscopatum, A. otavicum (Cockerell), A. polyochrum

Mavromoustakis, A. spilotum (Cockerell) = A. Kimberleyana (Friese), A.

sternale, A. tegwaniense (Cockerell), A. transversale Pasteels, A. zebra

(Friese) = Anthidiellum medionigrum Cockerell.

159

Diagnosis


2.5-5.0 mm; body: 3.5-6.0 mm. Vestiture: white, sparse to impubescent,

moderately long and dense around antennal sockets; male sternal comb on

apical region of S5; thick stout setae sometimes on tibiae; upper edge of inner

surface of hind tibia evidently marked by limited area of keirotrichiate.

Integument: black with yellow maculation on face, dorsal region of

preoccipital area, sometimes on margins of scutum, axilla and scutellum,

meso-lateral regions of metasoma; preoccipital area carinate dorsally;

paraoccular carina very reduced; vertex convex, 2X ocellus diameters above

eye; distance between antennal sockets more than two antennal socket

diameters; subantennal suture weakly curved, nearly 0.6X clypeus length;

scape long (7X first flagellar segment); antennal flagellum short (subequal to

head length); apical region of female mandible twice as broad as base;

pronotal carina small mesally, becoming progressively lamellate distally;

tegula narrow anteriorly, lateral margin slightly concave, except in A.

krugerense where lateral region of scutum becomes strongly carinate (Fig.

102); omaular weakly carinate, lamellate laterally, extends ventrally to midline

(left and right omaular join); axilla not extending laterally beyond lateral

margin of scutum; rows of pits across upper margin of propoduem very

reduced, evident only laterally, not carinate; T2-T5 of females, and to T6 of

males, in dorsolateral view, gently curved towards apex; propodeum

scrobiculate dorsolaterally; female T1 carinate antero-dorsally; T2-T5, in

dorsolateral view, curved gently downwards

160

Anthidiellum (Pycnanthidium) absonulum (Cockerell)

Anthidium absonulum Cockerell, 1932b: 526-527, male holotype (NHML)

Democratic Republic of the Congo; Mavromoustakis, 1937a: 232;

Mavromoustakis, 1937b: 267; Mavromoustakis, 1963b: 499.

Anthidium absonulum Cockerell [!]: Mavromoustakis, 1934: 43;

Mavromoustakis, 1963b: 499.

Anthidiellum absonulum (Cockerell): Cockerell, 1936c: 455.

Anthidiellum (Pycnanthidium) absonulum (Cockerell): Pasteels, 1984: 96, 99-

100.

Anthidiellum aspidopodium Mavromoustakis, 1936a: 45-47, female holotype

(NMBZ or NHML) Zimbabwe.

Pygnanthidium (Pygnanthidium) absonulum (Cockerell): Mavromoustakis,

1963b: 492.

Pygnanthidium (Pygnanthidium [!]) aspidopodium (Mavromoustakis):

Mavromoustakis, 1963b: 492; Pasteels, 1984: 99, syn.

Diagnosis


forewing 3.2-4.6 mm; body 4.5-5.9 mm. Vestiture: white; sparse, dense on

tibiae and tarsi; thick stout setae on tibiae. Integument: black, except yellow

maculation mesad to antennal socket, below antennal sockets, lower

paraocular area, most of clypeus (ventral clypeal margin black, Figure 24a), a

spot on dorso-lateral margin of preoccipital area, proximal end of dorsal

surface of fore tibia, dorsal surface of hind tibia (varying from basal spot to

161

almost entire dorsal surface, Figure 24g), meso-lateral regions of T1-T4 (wider

laterally on T3, medially interrupted cross-band on T4), continuous cross-band

on T5, almost entire T6 (Figure 24d); distal tarsal segments orange; mandible

punctate basally, finely sculptured apically, shiny between punctures, with

inner and outer carinae, two large outer teeth, two small inner teeth; lower

clypeal margin with four very small papillae (two on either side of mid-line);

lateral margin of scutum, in dorsal view, gently rounded; scuto-scutellar suture

open, forming shiny fovea; scutellum, in dorsal view, emarginate medio-

posteriorly, slightly obscures propodeum (Figure24c); 11 hamuli on hind

wing; vein cu-v straight; hind basitarsus broad, much broader than middle

basitarsus, long, half as long as hind tibia.

Male. Length: head 1.5-1.8 mm; scutum and scutellum 1.6-2.0 mm;

forewing 4.5-4.8 mm; body 5.2 mm. Vestiture and integument: similar to

female, except differ in the following characters: S5 with distal comb,

narrowly interrupted medially, about 28 similar setae; yellow maculation on:

entire clypeus, most of mandible, apex black (Figure 24b), ventral surfaces of

fore and middle femura, dorsal surfaces of fore and middle tibiae and basitarsi,

proximal and distal ends of dorsal surface of hind tibia, hind basitarsus (Figure

24f), medially interrupted cross-band on T3, continuous or medially

interrupted cross-band on T4, most of T5-T7, distal margins reddish; apex of

mandible half as wide as female, three subequal teeth (Figure 24b); S3 with

short, median notch; gonostylus expanded apically, slightly notched on outer

margin; penis valves seperated narrowly (Figure 24e).

162

Distribution

Anthidiellum absonulum is widespread throughout central, eastern and

southern regions of Africa.

163

Fig A1 Fig. A2

A. absonulum (female)

A. bipectinatum (female)A. absonulum

A. absonulum

A. absonulum

(male)

Figure 24a, b, c, d, e, f and g. Diagnostic features of Anthidiellum absonulum

(Cockerell). (a and b) Face of the female and male. (c and d) Mesosoma and

metasoma in dorsal view, arrows indicating regions of yellow maculations of

the female. (e) Male genitalia. (f and g) Hind legs of male and female

respectively.

A. a

bso

nulu

m

A. a

bso

nu

lum

(ma

le)

g

d

a

f

c

b

e

164

Anthidiellum (Pycnanthidium) apicatum (Smith)

Anthidium apicatum Smith, 1879: 84, female holotype (NHML) South Africa.

Anthidium nigripes Friese, 1904b: 105, female holotype (ZMHB)

Mozambique; Friese 1905b: 67; Friese 1909a: 400, 409; Friese 1909b:

158, 166; Strand 1912a: 143; Strand 1912b: 307; Pasteels 1984: 98,

syn.

Anthidium (Pygnanthidiellum [!]) apicatum Smith: Friese 1905b: 66; Friese

1909a: 400, 404; Friese 1909b: 158.

Anthidiellum (Pygnanthidiellum [!]) apicatum (Smith): Pasteels 1984: 98, 165.

Dianthidium melanocephalum Cockerell, 1920: 298-299, 300, three female

syntypes (NHML) South Africa; Pasteels 1984: 98, syn.

Anthidiellum nigripes var. rhodesinum Mavromoustakis, 1937a: 231, female

holotype (TMSA) Zimbabwe.

Anthidiellum nigripes var. rhodesianum [!] Mavromoustakis: Anonymous

1958: 33.

Pygnanthidium (Pygnanthidiellum) apicatum (Smith): Mavromoustakis

1963a: 492.

Diagnosis

Male. Unknown. Female. Length: head 1.7-2.4 mm; scutum and

scutellum 1.6-2.0 mm; forewing 3.7-4.4 mm; body 4.8-5.7 mm. Vestiture:

white; sparse; thick stout setae absent on tibiae. Integument: black, tegula, legs

Figure 25b) and metasoma dark brown, except yellow maculations on meso-

lateral regions of T1-T3 (yellow area increases progressively from a lateral

165

spot on T1 to occupying lateral third of T3), antero-lateral region of T4

yellow, medial regions of T5-T6 (Figure 25a); mandible with large basally,

small punctures apically, shiny between punctures, outer and inner carinae,

two large outer teeth and two similar inner teeth subequal to outer teeth;

clypeus margin with four projections, two on each side; lateral region of

scutum, in dorsal view, rounded; scuto-scutellar suture open, forming shiny

fovea; scutellum, in dorsal view, emarginate medio-posteriorly, hides median

region of propodeum; 11 long hamuli on hind wing; vein cu-v straight; hind

basitarsus broad, nearly as wide as base of hind tibia.

Anthidiellum (Pycnanthidium) auriscopatum (Strand)

Anthidium auriscopatum Strand, 1912b: 308, two female holotypes (HUMB)

Equatorial Guinea; Strand, 1920: 102.

Anthidiellum auriscopatum (Strand): Mavromoustakis, 1936a: 44, 45;

Pasteels, 1984: 99, syn.

Diagnosis

Male. Unknown. Female. Length: head 1.9-2.1 mm; scutum and

scutellum: 1.9-2.1 mm; forewing 3.7-3.9 mm; body 5.4-6.1 mm. Vestiture:

mostly white, except black to reddish on fore tibia; sparse; tibiae without thick

stout setae. Integument: black, brownish on legs, except yellow to reddish-

brown maculations on most of lower half of face (mandible mostly black,

Plate 23a), lateral region of axilla, dorso-lateral region of scutellum, basi-

dorsal and distal regions of tibiae (Plate 23b), entire surface of hind basitarsus

(figure 26b), meso-lateral regions of T1-T2 (yellow area increases

166

progressively from a lateral spot on T1 to occupying lateral third of T2); entire

anterior regions of T3-T5, T6 entirely yellow; mandible with large punctures

basally, small punctures apically, shiny between punctures, with outer carina

only, two, similar, large outer teeth and two, similarly small teeth; clypeus

margin with six small projections (one on median, two on lateral (Figure 26a);

lateral region of scutum rounded; scuto-scutellar suture open, forming shiny

fovea; scutellum weakly emarginated medio-posteriorly; 12 small hamuli on

hind wing, vein cu-v straight; hind basitarsus broad, as wide as base of hind

tibia.

Distribution

Anthidiellum auriscopatum is endemic to Congo and Equatorial

Guinea.

167

Figure 25a and b. Diagnostic features of Anthidiellum apicatum (Smith). (a)

Mesosoma, with arrow showing regions of yellow maculation. (b) Hind leg

A. auriscopatum (Female)

A. auriscopatum

Figure 26a and b. Diagnostic features of Anthidiellum auriscopatum (Strand).

(a) Face and (b) Hind leg, arrows showing regions of yellow maculations.

a b

Fig.

27

a

b

168

Plate 23a and b. Diagnostic features of Anthidiellum auriscopatum (Strand).

(a) Face and (b) Hind leg of with arrows indicating regions of yellow

maculation.

Plate 24a and b. Anthidiellum bipectinatum Pasteels. (a) Adult Female in

dorsolateral view. (b) Arrow showing regions of yellow integumental

maculation on the face.

c

d

a

b

169

Anthidiellum (Pycnanthidium) bipectinatum Pasteels

Anthidiellum (Pygnanthidiellum) bipectinatum Pasteels, 1984: 97, 105-106,

male holotype (AMNH); South Africa.

Diagnosis


forewing 3.9-4.7 mm; body 5.3-6.3 mm. Vestiture: white, sparse to

impubescent; tibiae without thick stout setae. Integument: black to dark brown

(lighter on tegula, legs and metasoma), with yellow maculations on most of

lower half of face (mandible and ventral margin of clypeus black, Figure 27a,

Plate 24a and b), proximal end of fore tibia, spot near base of middle tibia,

most of dorsal surface of hind tibia, entire dorsal surface of hind basitarsus

(Figure 27d), meso-lateral regions of T1-T4 (yellow area increases

progressively from a lateral spot on T1 to occupying lateral third of T4); entire

T5-T6 yellow (Figure 27c); mandible with large punctures at base and small

punctures towards apex, shiny between punctures, with both outer and inner

carinae, two, similar, large outer teeth and two, similarly small teeth; lower

clypeal margin with three projections, a median and two lateral; lateral margin

of scutum, in dorsal view, rounded; scuto-scutellar suture forms shiny fovea;

scutellum, in dorsal view, emarginate medio posteriorly, slightly obscures

propodeum; 11 small hamuli on hind wing, vein cu-v slants; hind basitarsus

broad, nearly as broad as base of hind tibia.


forewing 3.9-4.7 mm; body 5.8-6.3 mm. Vestiture and integument: similar to

170

female, except for gender limited characters and S3-S5 with short, distal

fringes; S5 with lateral comb widely seperated medially, with about eighteen

setae; clypeus entirely yellow (without black ventral margin), most of

mandible yellow, apex black (Figure 27b), ventral surface of fore femur,

dorsal surface of fore tibia, part of dorsal surfaces of middle and hind tibiae,

entire dorsal surfaces of basitarsi, dorsolaterally on T3, dorsally on T4, most

of T5-T7, distal margins reddish; apex of mandible half as wide as female,

teeth very small.

Distribution

Anthidiellum bipectinatum is endemic to southern Africa.

171

A. absonulum (female)

A. bipectinatum (female)

Figure 27a and b. Two diagnostic features of Anthidiellum bipectinatum. (a)

Face of the female and male arrows showing regions of yellow integumental

maculation. (b) Metasoma and (c) hind leg of the female, indicating regions of

yellow maculations.

A. bipectinatum(female)

a b

c

d

172

Anthidiellum (Pycnanthidium) bulawayense Mavromoustakis

Anthidiellum bulawayense Mavromoustakis, 1937a: 231-232, male holotype

(TMSA) Zimbabwe.

Anthidiellum (Pygnanthidiellum) bulawayense Mavromoustakis: Pasteels,

1984: 96, 99.

Diagnosis

Female. Unknown. Male. Length: head 1.3-1.5 mm; scutum and


white; sparse to impubescent, S2-S3 with short fringe on distal margin; S4

with long fringe, S5 with a lateral comb, (widely seperated medially) with

about fourteen subequal setae, thick stout setae absent on tibiae. Integument:

black, extremities reddish-black, yellow maculation on most of lower half of

face, black on dorso-medial region of supraclypeus (Figure 28), small yellow

spot on vertex, part of pronotal lobe and medio-lateral region of scutum,

narrow postero-lateral margin of scutellum, entire or distal part of ventral

surface of fore femur, dorsal surface of fore tibia, proximal and distal ends of

middle and hind tibiae, entire dorsal surface of middle and hind basitarsi,

meso-lateral region of T1 (small lateral spot), T2 similar to T1 or black, T3

similar to T4 or black, T4 (small yellow area in middle of lateral half),

posterior margin of T5 yellow, entire surfaces of T6-T7 (Plate 25); mandible

with large punctures basally and small punctures apically, shiny between

punctures, outer carina only, 1 large outer tooth, 2 similar, small inner teeth

(Figure 27); lower clypeal margin with 3-5 projections, one median, one-two

173

lateral; lateral margin of scutum, in dorsal view, rounded; scuto-scutellar

suture open, forming shiny fovea; scutellum, in dorsal view, emarginate

medio-posteriorly, hides median region of propodeum; 10 small hamuli on

hind wing; vein cu-v straight; hind basitarsus slender, half as long as hind

tibia.

Distribution

Anthidiellum bulaweyense is known from tropical Africa and dry

savannah in Zimbabwe.

Anthidiellum (Pycnanthidium) frontoreticulatum sp. nov.

Anthidiellum (Pycnanthidium) frontoreticulatum, female holotype (MRAC)

Kenya.

Description

Female. Length: head 1.5-1.6 mm; scutum and scutellum: 1.5-1.8 mm;

forewing 3.5-3 8 mm; body 4.8-5.0 mm. Vestiture: mostly white, brownish on

tarsi; sparse to impubescent, moderately dense around antennal socket,

mesopleuron, tibiae and tarsi; tibiae without thick stout setae. Integument:

mostly black, except yellow maculations on lower region of face (mandible

entirely black, Plate 26a), lateral regions of T1-T2, meso-lateral regions of T3-

T4 (small on T3), medial region of T6 (Figure 29a); mandible with large

punctures basally, small punctures apically, shiny between punctures, with

outer carina only, two, similar, large outer teeth and two, similarly small teeth;

174

clypeus margin with four small projections (two on each side); lateral region

of scutum rounded; scuto-scutellar suture open, forming shiny fovea;

scutellum, in dorsal view, truncate posteriorly; 11 small hamuli on hind wing,

vein cu-v slants; hind basitarsus broad, nearly as wide as base of hind tibia

(Figure 29c).


forewing 3.9-4.1 mm; body 4.2-4.5 mm. Vestiture: similar to female, except

for gender limited characters and S5 with lateral comb narrowly seperated

medially, with about 40 similar setae (c.f. Fig. 8). Integument: similar to

female, except clypeus entirely yellow (without black ventral margin), most of

mandible yellow, apex black, basal region of dorsal surfaces of fore, middle

and tibia, lateral region of T1; apex of mandible half as wide as female, teeth

very small; gonostylus apically expanded, notched on outer margin; penis

valves narrow apically, widely separated (Figure 29b)

Distribution

Anthidiellum frontoreticulatum sp. nov is endemic to Kenya.

175

A. bulaweyense(male)

Figure 28. Face of Anthidiellum bulawayense Mavromoustakis with arrows

showing regions of yellow maculation

A. frontorecticulumA. frontoreticulum

A. frontorecticulum

Figure 29a, b and c. Diagnostic features of Anthidiellum frontoreticulatum sp.

nov. (a) Metasoma, arrows showing regions of yellow maculations. (b) Male

genitalia. (c) Hind leg, arrow showing region of yellow maculation.

a

c

b

176

Plate 25. Male of Anthidiellum bulawayense Mavromoustakis.

Plate 26a, b and c. Diagnostic features two species of the genus Anthidiellum.

(a) Face of Anthidiellum frontoreticulatum sp. nov. with arrow showing

regions of yellow maculations. (b and c) Face and hind leg (b) of Anthidiellum

kimberleyana (Friese).

a

b

c

a

177

Anthidiellum (Pycnanthidium) kimberleyana (Friese)

Anthidium kimberleyana Friese, 1922: 30-31, female holotype (HUMB) South

Africa; Cockerell, 1936a: 3.

Anthidiellum kimberleyana (Friese): Mavromoustakis, 1936a: 44-45.

Anthidiellum (Pygnanthidiellum) kimberleyana (Friese): Pasteels, 1969: 46-

47; Gess, 1981: 25, 61; Pasteels, 1984: 106.

Diagnosis

Male. Unknown. Female. Length: head 1.9 mm; scutum and scutellum

2.0 mm; forewing 4.5 mm; body 5.0 mm. Vestiture: white, except reddish to

black on tarsi; sparse, tibiae without thick, stout setae. Integument: black, legs

reddish-brown to black (Figure 30c; Plate 26c), except yellow maculations on

paraocular area (Figure 30a; Plate 26b), a stripe on dorsal margin of

preoccipital area, antero-laterally on T1-T2, meso-laterally on T3-T4, T5 with

small yellow spots dorso-laterally (Figure 30b); mandible with large punctures

basally, shiny between punctures, with two carinae (outer and inner carinae),

two large dissimilar outer teeth and two small dissimilar inner teeth; lower

clypeal margin with five very small papillae (one median, two lateral); lateral

margin of scutum, in dorsal view, rounded; scuto-scutellar suture open,

forming shiny fovea; scutellum, in dorsal view, emarginated medio-

posteriorly, hides median region of propodeum; 12 hamuli on hind wing; vein

cu-v straight; hind basitarsi nearly as long as broad.

DISTRIBUTION

Anthidiellum kimberleyana (Friese) is endemic to South Africa.

178

A. kimberlyana(female) A. kimberlyana

A. kimberlyana (female)

Figure 30a, b and c. Diagnostic features of Anthidiellum kimberleyana

(Friese). (a) Face, arrows showing regions of yellow maculations. (b and c)

Metasoma and Hind leg of Anthidiellum kimberleyana (Friese) with arrows

indicating regions of yellow markings.

Fig

. 71 a

c

b

179

Anthidiellum (Pycnanthidium) madli Pauly

Anthidiellum (Pycnanthidium) madli Pauly [In] Pauly et. al., 2001: 229, male

holotype (NHMV) Madagascar.

Comment

This genus endemic to Madagascar was not available for examination

during this study, due to the fact that, it is a rare specimen unwilling to loan

them out.

Anthidiellum (Pycnanthidium) mediale Pasteels

Anthidiellum (Pygnanthidiellum) mediale Pasteels, 1984: 96, 101-103, male

holotype (EC) South Africa.

Comment

This genus was not available for examination during this study, due to

the fact that, it is a rare specimen and the museum was unwilling to loan then

out.

Anthidiellum (Pycnanthidium) micheneri Pauly

Anthidiellum micheneri Pauly [In] Pauly et al., 2001: 226, 228-229, female

holotype (SEMC) Madagascar.

180

Comment



out.

Anthidiellum (Pycnanthidium) orichalciscopatum (Strand)

Anthidium orichalciscopatum Strand, 1912b: 307-308, female holotype

(ZMHB) Equatorial Guinea.

Anthidiellum orichalciscopatum (Strand): Mavromoustakis, 1936a: 44.

Anthidiellum (Pygnanthidiellum) orichalciscopatum (Strand): Pasteels, 1984:

99.

Diagnosis

Male: Unknown. Female. Length: head 1.9-2.2 mm; scutum and


white, except brownish on tarsi; sparse to impubescent, moderate around

antennal socket and on mesopleuron; tibiae without thick, stout setae.

Integument: black, except extensive yellow maculation on most of lower half

of face, a spot on dorso-lateral margin of preoccipital area (Figure 31a), lateral

regions of scutum, axilla, posterior margin of scutellum, dorsal surface of fore

tibia, base of dorsal surface of middle tibia, hind mostly black (Plate 28),

meso-lateral regions of T1-T2, dorso-lateral and lateral regions of T3, short

continuous cross-bands on T4-T5 (Figure 31b; Plate 27a); mandible with large

punctures basally, small punctures apically, shiny between punctures, two

181

carinae (outer and inner carinae), two, similar, large outer teeth and two,

similarly small teeth; lower clypeal margin with four small papillae (two on

each side); lateral margin of scutum, in dorsal view, rounded; scuto-scutellar

suture open, forming shiny fovea; scutellum, in dorsal view, emarginate

medio-posteriorly, hides median region of propodeum; 12 hamuli on hind

wing; vein cu-v slants; hind basitarsus broad, nearly as wide as hind tibia.

Distribution

This species is widely distributed from central Africa to the southern

Cape of the coast of Africa.

182

Figure 31a and b. Two diagnostic features of Anthidiellum orichalciscopatum

(Strand). (a) Face and (b) Metasoma, with arrows indicating regions of yellow

maculation.

a b

Fig.

72

183

Plate 27a, b and c. Three species of genus Anthidiellum. (a) Anthidiellum

orichalciscopatum (Strand). (b) Anthidiellum otavicum (Cockerell). (c)

Anthidiellum polyochrum Mavromoustakis.

Plate 28. Hind leg of Anthidiellum orichalciscopatum (Strand), showing

reduced integumental markings.

a

b

28

c

184

Anthidiellum (Pycnanthidium) otavicum (Cockerell).

Dianthidium otavicum Cockerell, 1936c: 452-453, male holotype (NHML)

Namibia.

Anthidiellum (Pygnanthidiellum) otavicum (Cockerell): Pasteels, 1984: 96,

101, 103.

Diagnosis

Female. Length: head 1.5-1.6 mm; scutum and scutellum: 1.8-2.0 mm;

forewing 3.5-3.8 mm; body 4.5-4.7 mm. Vestiture: white, except black to

reddish on fore tibia; sparse; tibiae without thick stout setae. Integument:

black, except for yellow maculations on lateral and medio-dorsal regions of

clypeus, ventral region of paraocular area (Figure 32a), meso-lateral third of

T3-T5 (yellow spots on T3 smaller than on T4-T5, Plate 27b); mandible with

large punctures basally, small punctures apically, shiny between punctures,

with outer carina only, two, similar, large outer teeth and two, similarly small

teeth; lower clypeal margin with four small projections (two on each side

(Figure 32a)); lateral region of scutum rounded; scuto-scutellar suture open,

forming shiny fovea; scutellum weakly emarginate medio-posteriorly; 10

small hamuli on hind wing, vein cu-v straight; hind basitarsus broad, as wide

as base of hind tibia.

Male. Length: head 1.5-1.7 mm; scutum and scutellum: 1.6-1.7 mm;


female, except gender limited characters and S3-S5 with short, distal fringe;

S5 with a lateral comb, with about 20 similar setae, seperated narrowly by

Fig.

72

185

medial notch (Figure 4b); clypeus entirely yellow, ventro-lateral regions of

supraclypeus, mandible yellow with apex black; ventral surfaces of fore and

middle femora; dorsal surfaces of fore, middle and hind tibiae, dorsal surfaces

of basitarsi, lateral regions of T1-T3; cross band on T4 interrupted medially by

narrow blackish area; continuous cross band; T5-T7 mostly yellow; apex of

mandible half as wide as female, teeth very small; S5 with a small medial

notch; gonostylus expanded apically, margin with reduced notch; penis valve

narrowly seperated (Figure 32b).

Distribution

Anthidiellum otavicum (Cockerell) is endemic to south- western

Africa.

186

A. otavicum A. otavicum

Figure 32. Diagnostic features of Anthidiellum otavicum (Cockerell). (a) Face

and (b) Male genitalia, arrows indicate regions of yellow markings.

A. polyogrum

Figure 33. Hind leg of Anthidiellum polyochrum Mavromoustakis (arrows

showing regions of yellow markings)

a b

33

187

Anthidiellum (Pycnanthidium) polyochrum Mavromoustakis

Anthidiellum polyochrum Mavromoustakis, 1937b: 266-267, female holotype

(SAMC)

Anthidiellum (Pygnanthidiellum) polyochrum Mavromoustakis: Pasteels,

1984: 97, 102-103, 105.

Diagnosis


forewing 3.4-3.5 mm; body 3.8-4.2mm. Vestiture: white; sparse to

impubescent, except moderately dense around antennal socket and on tarsi;

tibiae without thick stout setae. Integument: mostly black, except yellow

maculation on most of lower face, basal region of mandible, dorsal region of

preoccipital area, dorsal surfaces of fore tibia and basitarsus, basal part of

dorsal surfaces of middle, hind tibia and most of hind basitarsus (Figure 33),

meso-lateral regions of T1-T2, T3 with small lateral spot and larger dorso-

lateral spot, T4-T5 with dorso-lateral spot, T6 mostly yellow (Plate 27c);

mandible with large punctures basally, shiny between punctures, outer and

inner carinae, 2 large outer tooth, 2 similar, small inner teeth; lower clypeal

margin with 3-5 projections, one median, one-two lateral, lateral margin of



region of propodeum; 11 hamuli on hind wing; vein cu-v slants; hind

basitarsus broad, subequal to width of hind tibia.

188

Male. Length: head 1.3 mm; scutum 1.6 mm; forewing 2.5 mm; body

3.5 mm. Vestiture and integument: similar to female, except for sex limited

characters and S5 with lateral comb with 30 similar setae; apex of mandible

half as wide as in female, teeth very small.

Distribution

Anthidiellum polyochrum Mavromoustakis is endemic to Namibia.

Anthidiellum (Pycnanthidium) spilotum (Cockerell)

Dianthidium spilotum Cockerell, 1920: 299, female holotype (NHML) South

Africa; Cockerell, 1936a: 1, 6.

Anthidiellum (Pyganthidiellum) spilotum (Cockerell): Pasteels, 1984: 97, 106-

108.

Anthidium capuzinum Friese, 1925: 510-511; Pasteels, 1984: 106, syn.

Diagnosis


forewing 4.3-4.9 mm; body 5.6-5.9 mm. Vestiture: white, except reddish-

black on dorsal surface of tibiae; sparse to impubescent; thick stout setae

absent on tibiae. Integument: black, tegula and legs dark brown to black,

yellow maculation on ventro-lateral region of clypeus, basal region of

mandible (Figure 34a), most of paraocular area, dorso-lateral region of

preoccipital area, lateral region of scutellum, most of dorsal surfaces of fore,

middle and hind femora, antero-dorsal region of fore and middle tibiae,

189

proximal and distal ends of hind tibia, entire dorsal surfaces of fore and hind

basitarsus (Figure 34g; Plate 30a), meso-lateral regions of T1-T5 (narrow on

T1-T2, wide on T3-T5), medial region of T6 (Figure 34a; Plate 29); mandible

with large punctures basally, small punctures apically, shiny between

punctures, two carinae (outer and inner carinae), four distinct teeth (outer teeth

largest, third and fourth progressively smaller Figure 34a); lower clypeal

margin with six projections, one median, two on each side; lateral region of



region of propodeum; 11 long hamuli on hind wing; vein cu-v slants; hind

basitarsus broad, more than half as wide as hind tibia.

Male.Length: head 1.5-1.6 mm; mesosoma 1.5-1.7 mm; forewing 4.4-

4.9 mm; body 4.8-5.6 mm. Vestiture and integument: similar to female, except

gender limited characters and S2-S4 with short fringes; S5 with a lateral comb

with about 10 similar setae (latertal combs widely seperated medially;

scutellum and axilla entirely black, legs brownish, yellow maculation on most

of lower half of face (without black ventral margin), most of mandible (black

apically (Figure 34b)), anterior and distal regions of femora, basal regions of

fore and middle tibiae, proximal and distal regions of hind tibia (Figure 34f),

entire surface tarsi, wider on dorso-lateral region of T4, entire anterior region

of T6 (Figure 34d); apex of mandible half as wide, teeth very small;

gonostylus expanded apically, slightly notched on outer margin; penis valves

widely separated (Figure 34e).

190

Distribution

Anthidiellum spilotum Cockerell is endemic to south-western and

southern Africa.

191

Figure 34a, b, c, d, e, f and g. Diagnostic features of Anthidiellum spilotum

(Cockerell). (a and b) Face of the female and male, arrows indicating areas of

yellow integumental maculations. (c and d) Mesosoma, of the male and female

respectively, arrows indicating area of yellow maculations. (e) Male genitalia.

(f and g) Hind legs of the male and female, respectively.

Fig

.

78

a b c

g

d e

f

F

i

g

.

7

8

Fig. 81

Fig.

81 F

i

g

.

8

1

192

Plate 29. Female Anthidiellum spilotum (Cockerell).

Plate 30a and b. Two diagnostic features of genus Anthidiellum. (a) Hind leg

of A. spilotum (Cockerell), arrows indicating regions of yellow maculations.

(b) Male genitalia of Anthidiellum sternale Pasteels.

a

b

29

193

Anthidiellum (Pycnanthidium) sternale Pasteels

Anthidiellum (Pygnanthidiellum[!]) sternale Pasteels, 1984: 96, 104-105, 155,

male holotype (TMSA) South Africa.

Diagnosis

Female. Length: head 1.7 mm; scutum and scutellum 1.8 mm;

forewing 4.4 mm; body 5.9 mm. Vestiture: white; sparse to impubescent; thick

stout setae on tibiae. Integument: black, antennal flagella and tegula dark

brownish, except yellow maculation on most of clypeus (dorsal and ventral

margins black), lateral region of supraclypeal area, lower half of paraocular

area (Figure 35a), dorsal region of preoccipital area, most of surfaces of

femora, entire tibiae and tarsi (Figure 35g), meso-lateral regions of T1-T2,

dorso-lateral regions of T3-T4 (mediolongitudinal black areas broad on T3,

narrow on T4), entire T5-T6 (Figure 35c); mandible with large punctures

basally, small punctures apically, shiny between punctures, with two carinae

(outer and inner carinae), two, similar, large outer teeth and two, similarly

small teeth; lower clypeal margin with four projections (two on each side);

lateral region of sutum, in dorsal view, rounded; scuto-scutellar suture open,

forming shiny fovea; scutellum, in dorsal view, emarginate medio-posteriorly,

hides median region of propodeum; 11 long hamuli on hind wing; vein cu-v

slants; hind basitarsus broad, more than half as wide as hind tibia.



female, except sex limited character and S5 with lateral comb of about 12

194

similar setae (combs widely seperated medially); apex of mandible half as

wide as in female, 3 very small teeth (Figure 35b), medial region of hind tibia

(Figure 35f); lateral region of T5 (Figure 35d); gonostylus slender apically,

deeply notched on outer margin; penis valves slightly wide apically, narrowly

separated (Figure 35e; Plate 30b).

Distribution

This species is endemic to South Africa.

195

Figure 35a, b, c, d, f and g. Diagnostic features of Anthidiellum sternale

Pasteels. (a and b) Face and mesosoma of the female and male, indicating area

of yellow maculations. (c and d) Metasoma of the female and male with

arrows showing regions of yellow markings. (e) Male genitalia. (f and g) Hind

legs of the male and female, respectively (arrow showing region of yellow

marking).

e

a a b

c d

f

F

i

g

.

7

6

Fig. 76

g

196

Anthidiellum (Pycnanthidium) tegwaniense (Cockerell)

Dianthidium tegwaniense Cockerell, 1914a: 278, female holotype (NHML)

South Africa.

Anthidiellum tegwaniense (Cockerell): Cockerell, 1932c: 524; Cockerell,

1936c: 455; Mavromoustakis, 1936a: 46.

Anthidiellum (Pygnanthidiellum) tegwaniense (Cockerell): Pasteels, 1984:

107-108.

Anthidiellum swalei Mavromoustakis, 1936b: 603-605, female holotype

(NHML) Zimbabwe; Pasteels, 1984: 107-108, syn.

Pygnanthidiellum (Pygnanthidiellum) swalei (Mavromoustakis):

Mavromoustakis, 1963b: 492-493.

Diagnosis


forewing 4.3-4.6 mm; body 5.4-5.8 mm. Vestiture: white; sparse to

impubescent; tibiae without thick stout setae. Integument: black, antenna,

tegula and legs dark brown, yellow maculations on most of lower half of face

(ventral margin of clypeus dark brown), basal region of mandible (apex

black), dorso-lateral region of preoccipital area, lateral regions of scutum and

axilla, dorso-lateral region of scutellum, entire legs (Figure 36d), meso-lateral

regions of T1-T4 (smaller on T1-T2, wider on dorso-lateral regions of T3-T4),

entire surfaces of T5-T6 (Figure 36b); mandible with large punctures basally,

shiny between punctures, outer carina only, two, similar, large, outer teeth and

two, similarly, small teeth; lower clypeal margin with four projections (two on

each side; lateral region of

197


fovae; scutellum, in dorsal view, emarginate medio-posteriorly, hides median

region of propodeum; 12 long hamuli on hind wing; vein cu-v straight; hind

basitarsus broad, nearly as wide as base of hind tibia.

Male. Length: head 1.5-1.8 mm; mesosoma 1.6-1.8 mm; forewing 3.5-

3.7 mm; body 4.9-5.4 mm. Vestiture and integument: similar to female, except

for gender limited characters and S5 with 30 similar setae laterally; yellow

maculation on entire clypeus, mandible (apex black, figure 36a), dorsal

surface of fore tibia, medial region of hind tibia (Figure 36c), smaller on T3,

most of T4-T7; apex of mandible half as wide, 3 very small teeth; gonostylus

narrow apically, deeply notched on outer margin; penis valves expanded

apically, widely separated (Figure 36e).

Distribution

Anthidiellum tegwaniense (Cockerell) is widespread throughout Malawi,

South Africa and Zimbabwe.

198

A. tewaniense

Figure 36a, b, c, d and e. Anthidiellum tegwaniense (Cockerell), indicating

regions of yellow integumental maculations. (a) Face of the male. (b)

Metasoma of the female. (c and (d) Hind legs of the male and female,

respectively. (e) Male genitalia.

Fig.

76

a b

c d

e

199

Anthidiellum transversale Pasteels

Anthidiellum (Pygnanthidiellum) transversale Pasteels, 1984: 104, 106, male

holotype (RMNH) Zimbabwe.

Comment



out.

Anthidiellum (Pycnanthidium) zebra (Friese)

Anthidium zebra Friese, 1904b: 103, male and female syntypes (TMSA) South

Africa; Friese, 1905b: 67-68, 70, 73; Friese, 1909a: 400, 401, 402, 409,

412; Friese, 1909b: 158, 160-161; Strand, 1912a; 142; Cockerell,

1932b: 527; Cockerell, 1936a: 6; Anonymous, 1958: 33.

Dianthidium zebra (Friese): Cockerell, 1909b: 261; Cockerell, 1920b: 299;

Cockerell, 1936a: 6; Cockerell, 1936c: 453.

Anthidiellum zebra (Friese): Mavromoustakis, 1936b: 605; Mavromoustakis,

1937a: 232; Mavromoustakis, 1937b: 267;

Pygnanthidiellum (Pygnanthidiellum) zebra (Friese): Mavromoustakis, 1963a:

492-493.

Anthidiellum (Pygnanthidiellum) zebra (Friese): Pasteels, 1984: 95-99, 101,

104, 106.

Anthidiellum medionigrum Cockerell, 1933a: 464, male holotype (NHML)

South Africa; Cockerell, 1936c: 455; Mavromoustakis, 1936a: 46;

Pasteels, 1984: 97, syn.

200

Diagnosis


forewing 4.3-4.5 mm; body 3.9-4.8 mm. Vestiture: white; sparse to

impubescent; thick stout setae on tibiae. Integument: black, tegula, legs and

metasoma dark brown, yellow maculations on most of lower half of face, most

of mandible (ventral margin of clypeus dark brown, Figure 37a), dorsal region

of preoccipital area, lateral regions of scutum and axilla, dorso-lateral region

of scutellum, small on ventral and distal regions of fore and middle femur,

proximal regions dorsal surfaces of fore and middle tibia, most of dorsal

surface of hind tibia, entire hind basitarsus (Figure 37g), meso-lateral regions

of T1-T4 (yellow area increases progressively from a lateral spot on T1 to

occupying lateral third of T4), entire surfaces of T5-T6 (Figure 37c); mandible

with large punctures basally, shiny between punctures, with two carinae (outer

and inner carinae), two, dissimilar, large outer teeth and two, similarly small

teeth; lower clypeal margin with four projections, two on each side; lateral

region of scutum, in dorsal view, rounded; scuto-scutellar suture open,

forming shiny fovea; scutellum, in dorsal view, emarginate medio-posteriorly,

hides median region of propodeum; 11 small hamuli on hind wing, vein cu-v

straight; hind basitarsus broad, subequal base of hind tibia.



female, except for sex limited characters and S2 with lateral fringe, S3-S4 with

narrow fringes, S5 with lateral comb of about 30 setae; yellow maculation on

entire dorsal surface of fore tibia, medial region of hind tibia (Figure 37f);

201

smaller on T4, most of T5-T7 mostly yellow, distal margins of T5-T6 reddish

to dark brown (Figure 37d); apex of mandible half as wide, 3 very small teeth

(Figure 37b); gonostylus slender aically, deeply notched on outer margin;

penis valves widely separated (Figure 37e).

Distribution

This species is endemic to southernmost part of Eastern and South

Africa

202

Figure 37a, b, c, d, f and g. Diagnostic features of Anthidiellum zebra (Friese).

(a and b) Face of the female and male, arrows indicating regions of yellow

integumental maculations. (c and d) Metasoma of the female and the male

respectively (arrow indicate regions of yellow markings). (e) Male genitalia. (f

and g) Hind legs of the male and female, respectively.

a b c d

F

i

g

.

7

6

e

Fig. 76

Fig. 76

f

g

Fig. 76

203

Anthidiellum (Pycnanthidium) krugerense sp. nov.

Anthidiellum (Pycnanthidium) krugerense, female holotype (SANC), South

Africa.

Description

Female: Length: head 1.9 mm; scutum and scutellum 1.8 mm; forewing 4.0

mm; body: 4.8 mm. Vestiture: white; sparse to impubescent; thick stout setae

on tibiae. Integument: black, tegula, lateral regions of axilla and scutellum

dark brown, except for yellow maculations on most of lower half of face

(ventral clypeal margin black, Figure 38a), dorso-lateral region of preoccipital

area, entire legs, meso-lateral regions of T1-T4 (yellow area increases

progressively from a lateral spot on T1 to occupying lateral third of T4), entire

surfaces of T5-T6; mandible with small, distinct punctures, shiny between

punctures, outer carina only, two large similar outer teeth and two small

dissimlar inner teeth; lower clypeal margin with four projections, two on each

side; lateral margin of scutum, in dorsal view, extend longitudinally concave,

groove separate extended surface with carina next to tegula (Figure 38b);

scuto-scutellar suture open, forming shiny fovea; scutellum, in dorsal view,

emarginate medio-posteriorly, hides median region of propodeum; 12 long

hamuli on hind wing; vein cu-v straight; hind basitarsus broad, nearly as wide

as base of hind tibia.

Distribution

This species is endemic to South Africa.

204

Figure 38a and b. Descriptive features of Anthidiellum krugerense sp. nov. (a)

Face, arrows showing region of yellow markings. (b) Scutum, arrow

indicating the carina on the lateral region of scutum extending beyond axilla

margin.

b a

205

CHAPTER FIVE

DISCUSSION

Phylogenetic analysis of the tribe Anthidiini

The tribe Lithurgiini is reported as the sister group of the Osmiini,

Anthidiini, Megachiliini and Dioxyini which constitute the Megachilinae, with

the tribe Megachilini known to the closest sister group of the Anthidiini (Riog-

Alsina and Michener, 1993). These authors adequately demonstrated the

Megachilinae to be monophyletic by the presence of several unique

synapormorphies. However, they did not demonstrate the monophyly of the

Anthidiini using most of the genera of the tribe. The inclusion of Lithurge

pullatus (Vachal) and Megachile semiflava (Cockerell) in this study as two

outgroups clearly confirmed the monophyly of the Anthidiini. This is

supported by nine synapormorphies, seven of which are non-homoplasious

namely, (a) Lateral region of preocciptal area of female carinate (level of

sclerotization variable. (b) Distance between antennal sockets at least subequal

to two antennal sockets diameters. (c) Omaulus carinate (sometimes entirely)

(d) Scutellum, in dorsal view, obscures propodeum, at least medially. (e)

Vestiture on dorsal surface of hind basitarsus dense. (f) Second recurrent vein,

usually extends distally behind second transverse cubital vein. (g) Jugal lobe

short, less than half length of vannal lobe. (i) Anterior region of T1 without

fovea. (j) Metasoma without subapical hair bands. Characters (d) and (e) show

206

reversals in higher tree topology and therefore probably do not strictly

characterize the Anthidiini as a whole. The Anthidiini is clearly supported by

several unambiguous synapormorphies and by very high bootstrap and

jackknife values of 96% and 97% respectively, hence confirms the tribe as a

monophyletic taxon.

The second major island comprises relatively small genera, namely

Aspidosmia, Afranthidium, Anthidioma, Neanthidium, with very high

boothstrap and jackknife valves of 86% and 85% in the strict consensus trees.

Aspidosmia forms the first major clade within this island. Its monophyly is

demonstrated by an autoapormorphy, that is, the dorsal region of the female

clypeus being highly elevated. This uniquely derived autoapomorphy is

supported by another character state namely, second recurrent vein of the

forewing entering the second submarginal cell. The clade therefore supports

the relationship of genus Aspidosmia to the rest of the anthidine bees which

confirms previous classification works including Peter (1972), Riog-Alsina

and Michener (1993), Michener and Griswold (1994) and Michener (2000).

The genera Afranthidium, Anthidioma and Neanthidium formed the next sub-

clade within this island. The monophyly of this clade was demonstrated by

two synapomorphies: apex of marginal cell mostly rounded and the first

recurrent vein of the fore wing meeting the first transverse cubital vein distally

or entering the first submarginal cell. The high evidential support of this clade,

gives an indication of a very close relationship between these three genera.

However, in the absence of the male specimen of Anthidioma, examination of

the male genitalia could not be done to confirm the ascertion by Michener and

207

Griswold (1994) and Michener (2000) that this genus should be placed as a

subgenus of Afranthidium. Hence Afranthidium and Anthidioma have been

retained as separate genera. In addition, due to the considerable variations in

the number of spines on T6 and T7 of male, structure of the male genitalia and

in the mandibular dentition of females of Afranthidium and Neanthidium, the

two genera were also retained as separate anthidine taxa as had been

documented in Michener (2000).

Two large groups constitute the remaining major islands within the

Anthidiini. One of these comprises three parasitic genera and nine non-

parasitic. The parasitic genera include Afrostelis, Euaspis and Hoplostelis

while the non-parasitic genera comprise Anthidioctes, Hypanthidiodes,

Cyphanthidium, Paranthidium, Eoanthidium, Epanthidium, Serapista,

Duckeanthidium, Apianthidium and Aztecanthidium. This island is clearly

supported by high bootstrap and jackknife values of 83% and 82%

respectively and gives an indication that these genera form a close sister

group. In addition, two synapomorphies including a non-homplasious

character also support this island namely, the posterior surface of the antennal

scape being sparsely pubescent and the facial vestiture also being sparse to

impubescent. The latter reversed in higher tree topology, probably due to the

fact that, that character is commonly shared by other genera. In the analysis of

this island of trees, the clade [Anthidioctes, Hypanthidiodes, Afrostelis and

Hoplostelis] is not defined by any unique synapomorphy. However, Afrostelis

and Hoplostelis are shown clearly as monophyletic, defined primarily by the

elongated scutum and scuto-axilla suture slightly opening to form a fovea as

208

well as absence of a scopa of the female. These parasitic genera supported by

low bootstrap and jackknife values (51/ 50%), clearly show little support of

this group. On the other hand, Anthidioctes and Hypanthidiodes are defined by

high bootstrap and jackknife values (78/ 78%), hence have high evidential

support of this group. Synonymization of the Neotropic Anthidioctes with

Neotropic Hypanthidiodes is suggested in this present study. Cyphanthidium

forms the third sub-clade of this island. Inspite of its high bootstrap and

jackknife values (83/ 86%), the genus however is not defined by any

autapomorphy. Thus, in this study, the monophyly of this genus was not

established. The monophyly of genus Paranthidium, which constitute the fifth

sub- clade was not demonstrated in this study because of the absence of an

autapomorphy. However, the combination of two homoplasious character

states defined this clade, i.e. a lower clypeal margin with projections and a

slight notch in the scutellum postero-medially. This clade as sister group is

supported by very high bootstrap and jackknife values (91/ 90%). The clade

Euaspis, Eoanthidium and Epanthidium forms the sixth sub-clade within this

island. The parasitic genus Euaspis, is strongly supported by bootstrap and

jackknife values (90/ 90%) and is characterized primarily by the scutellum, in

dorsal view, being deeply notched postero-medially. Inspite of the presence of

two homoplasious synapomorpies in defining genus Eoanthidium as

monophyletic, it forms polytomy with Neotropic genus Epanthidium which is

not characterized by an autoapomorphy. These two genera are weakly

supported evidentially by low bootstrap and jackknife values 51/ 50%.

Inadequate information could probably account for the poor resolution of the

209

clade. The monophyly of genus Serapista is strongly supported by a single

homoplasious apomorphy inwhich the posterior region of the axilla has a

spine. The high bootstrap and jackknife values (98/ 98 %) support the clade

that includes Serapista and three other genera namely, Duckeanthidium,

Apianthidium and Aztecanthidium. The clade formed by these three genera is

primarily defined by a single synapomorphy inwhich the apex of the outer

hind tibia spur is strongly curved. Furthermore, a high bootstrap and jackknife

values 87/ 85% evidentially support this clade. The monophyly of

Duckeanthidium was defined by one homoplasious character state, i.e.

vestiture on the dorsal surfaces of tibiae and basitarsi is densely pubescent. In

addition, Apianthidium was defined by a single apomorphy inwhich the

second recurrent vein of the fore wing enters the second submarginal cell.

Aztecanthidium on the other hand was not characterized by any derived

character state, hence its monophyly was not established in this study.

The remaining major island of the Anthidiini comprised Anthidium,

Dianthidium, Anthidiellum, Bathanthidium, Icteranthidium, Gnathanthidium,

Pachyanthidium, Pseudoanthidium, Acedanthidium, Benanthis, Hypanthidium,

Indanthidium, Notanthidium, Pleisianthidium, Rhodanthidium and Trachusa.

This island is supported by high bootstrap and jackknife values (80/ 79 %) and

is also defined by a combination of two characters namely, the second cubital

cell of the fore wing is more than half or subequal length to the length of the

first cubital cell and also, vein Cu-v of the hind wing is slanted. The genus

Anthidium forms a clade within this island and is seperated from the rest by

two synapomorphies: (a) dorsal region of preoccipital area, in female carinate

210

and b) facial vestiture sparse to impubescence. High bootstrap and jackknife

values of 82/ 82% clearly support this clade. Anthidium was establisted as

monophyletic, defined by a single homoplasious character state in which the

second recurrent vein meets the second transverse cubital vein distally. The

next sub-clade within that island consists of the genera Dianthidium,

Anthidiellum and Bathanthidium. This clade is not defined by any

synapomorphy, although it is highly supported by bootstrap and jackknife

values of 63/ 63 %. The monophyly of Dianthidium is defined primarily the

female having three mandibular teeth. Anthidiellum and Bathanthidium

formed a subclade strongly supported by very high bootstrap and jackknife

(85/ 84%) and defined by two homoplasious synapomorphies, i.e. a curved

subantennal suture and foveate scuto-scutellar suture. In the absence of male

characters of Bathanthidium in this study, synonymization the two taxa are in

this study was not considered as was suggested in Michener (2000). The

monophyly of Anthidiellum was demonstrated by two homoplasious

apomorphies, i.e. projections on the ventral margin of the clypeus and a

slighty notch on the scutellum postero-medially. A single non-homoplasious

synapomorphy separated this clade from the rest, i.e. posterior margin of the

axilla extending beyond the lateral margin of the scutum. The genera

Gnathanthidium, Pachyanthidium and Pseudoanthidium formed the next sub-

clade, and were characteristically defined by a single non-homoplasious

synapomorphy in which the preoccipital area is lamellate. Low bootstrap and

jackknife values (50/ 50%) weakly supported this clade to be considered as

sister group. However, the subclade Gnathanthidium, Pachyanthidium and

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Pseudoanthidium was clearly supported by high bootstrap and jackknife

values (83/ 85%, and 79/ 79%). All three genera are defined by at least one

apomorphy, which demonstrates their monophyletic status. Icteranthidium

was however not defined by any uniquely derived character hence its

monophyly was not confirmed in this present study. This clade was separated

from the remaining clade by one non-homoplasious character state and two

homoplasious character states, as well as, high bootstrap and jackknife values

(81/ 80%) supporting this clade as sister group. The remaining clade which is

weakly supported by bootstrap and jackknife values (63/ 65%) had several

synapomorphies defining the monophyly of the genera. This clade comprises

Acedanthidium, Benanthis, Hypanthidium, Indanthidium, Notanthidium,

Pleisianthidium, Rhodanthidium and Trachusa. The collapse of several nodes

within this clade could probably be due to inadequate characters to clearly

resolve the clade. The genera Rhodanthidium, Hypanthidium and

Pleisianthidium were not supported by any autoapomorphy to ascertain their

monophyly. However, these genera differ significantly in the structure of the

male genitalia, hence, could be maintained as separate taxa.

In the analysis of the genus Anthidiellum, the first clade consists of

Anthidiellum flavescens, A. eritrinum and A. rubellum. The clade is

evidentially supported by very high bootstrap and jackknife values (94/ 95%)

and three synapomorphies namely, the distance between antennal sockets

subequal or less than two antennal socket diameters and a preoccipital carina

occurring dorso-laterally or laterally, as well as a distally lamellate pronotal

lobe. However, the two species of the nominative subgenus formed a sub-

212

clade and was highly supported by bootstrap and jackknife values of 73/ 73%.

Hence, this study clearly confirms the two clades as separate subgenera

(Anthidiellum and Chloranthidiellum) within the genus as in Michener (2000).

The second major clade is made of A. absonulum, A. apicatum, A.

auriscopatum, A. frontoreticulatum, A. kimberleyana, A. polyochrum, A.

krugerense, A. sternale, A. tegwaniense, A. zebra, A. bulawayense, A.

otavicum, A. bipectinatum and A. spilotum. The clade is supported by very

high bootstrap and jackknife values (92/ 89%), and two synapomorphies

inwhich the length of the subantennal suture is more than half the length of the

clypeus and mandibles having small punctures apically and large punctures

basally. Several nodes collapsed to form polytomies within this clade. The

reason for this could be due to the inavailability of several specimens of most

taxa to adequately resolve the cladogram.

In spite of a number of missing genera and character states during the

analysis of the tribe, at least the following classification, based on available

information, is postulated for the tribe. Most of the generic classification of

Michener (2000) is retained, except that, three main subtribes are erected

within the tribe. The genus Hypanthidiodes is synonymized with Anthidioctes

(the latter being the older available name). In addition, synonymization of the

genus Bathanthidium with genus Anthidiellum was considered, however, they

were retained as separate taxa. The tribe Anthidiini may therefore be placed

under family Megachilidae and subfamily Megachilinae with three subtribes,

namely Afranthidina, Anthidina and Euaspina. The subtribe Afranthidina

comprises non-parasitic, small to medium size, megachiliform to

213

anthophoriform and hairy bees which are all endemic to the African continent.

The four genera that occur in this subtribe include Afranthidium, Anthidioma,

Aspidosmia and Neanthidium. The Anthidina comprises mainly non-parasitic

bees with wide morphological variation. Its size ranges from medium to large

and varies in shape from hoplitiform, megachiliform, chalicodomiform to

anthophoriform. It is the only subtribe with three cosmopolitan genera

(Anthidium, Anthidiellum and Trachusa) while the other genera

(Acedanthidium, Bathanthidium, Benanthis, Dianthidium, Gnathanthidium,

Hypanthidium, Icteranthidium, Indanthidium, Notanthidium, Pachyanthidium,

Pleisianthidium and Pseudoanthidium) are known to have narrow range of

distribution. The subtribe Euaspina constitute both parasitic and non-parasitic

genera, just as in Anthidina, it varies from small to large size hoplitiform to

anthophoriform bees, as well as small heriadiform bees. The subtribe is made

up of genera with limited geographic distribution among the continents. These

genera include, Anthidioctes, Afrostelis, Apianthidium, Aztecanthidium,

Cyphanthidium, Duckeanthidium, Eoanthidium, Epanthidium, Euaspis,

Hoplostelis and Serapista. The classification of the Anthidiini into three

subtribes enormously facilitates its study and provides useful information on

these genera. The name of each subtribe was derived from the oldest available

name within the genera.

Biogeography and Evolution of Anthidiini

Based on the results of the present analyses, three main lineages of

Anthidiini are distinguished. The first lineage comprises a relatively small

clade of mostly Afrotropical genera (Afranthidium, Anthidioma and

214

Aspidosmia), and a Palaearctic genus Neanthidium (Algeria and Morocco).

The second lineage is made up of a distinctly large clade with three

cosmopolitan genera (Anthidium, Anthidiellum and Trachusa), three genera

endemic to the new world (Neotropic and Nearctic- Dianthidium,

Hypanthidium and Notanthidium), and ten genera occurring in the old world

(Afrotropical and Palaearctic: Icteranthidium, Pachyanthidium,

Gnathanthidium, Benanthis; Palaearctic and Orient: Rhodanthidium,

Bathanthidium, Acedanthidium, Indanthidium; Pleisanthidium is widespread

throughout the old world). The third clade also comprises a relatively large

clade of the remaining genera, with endemic biogeographical distribution;

eight genera of the new world (Neotropic and Nearctic: Anthidioctes,

Hoplostelis, Paranthidium, Aztecanthidium, Duckeanthidium), two

Afrotropical genera (Afrostelis, Cyphanthidium), one Oriental genus,

Apianthidium, and three genera with relatively extensive distribution within

the old world (Euaspis, Eoanthidium, Serapista).

These biogeographical data at the subtribe and generic levels were

used to obtain information about probable evolutionary processes and events

that might have taken place within the Anthidiini. The Afrotropical and

Oriental regions exhibit high bee richness within the Anthidiini at the generic

level, with 16 and 13 genera, respectively. One subtribe is limited to the Africa

continent. High bee richness at the generic level and basal groups of primitive

forms are enough evidence to postulate that, the origin and center of radiation

for the Anthidiini probably occurred in southern Pantropic Region (Southern

Africa to South East Asia). These geographic locations, before the split of the

215

world’s continent, formed South-eastern block of the southern hemisphere,

super-continent Gondwanaland (during the early Cretaceous over 100 million

years before present, Raven and Axelrod, 1974).

The discovery of the oldest fossil bee, Melittosphex burmensis

(Melittosphecidae) (Pionar and Danforth, 2006), from the early Cretaceous

Burmese amber (~100 million years ago) which is 35-45 million years older

than the previously known fossil bee, Cretotrigona prisca by Michener and

Grimaldi, 1988 a & b), tend to suggest that the evolution of bees might have

occurred earlier than previously indicated, possibly, as early as during the

evolution and diversification of angiosperms in early Cretaceous. M.

burmensis shares several characters with most members of present day

anthidine genera. Visual observation clearly shows that the forewing of this

bee possess strong first and second transverse cubital veins that forms two

complete submarginal cells. Further to this, this fossil resin-collecting bee has

a heriadiform shape which is typical of small size anthidine bees. Hence, it is

probable to also assume that the present day anthidine genera existed about the

same era as M. burmensis.

It is also known that, it was only in the tertiary (66.4 to 1.6 million

years ago) during the late Paleocene and early Eocene era that bilaterally

symmetrical organization of floral organs coevolved with animal behaviour

including bee sociality (Danforth, 2004; Dilcher, 2006). Based on the

discovery of the new fossil bee (after publication of Engel and Perkovsky,

2006), and data from host-plant specialization in western anthidine bees by

Müller (1996), as well as information of the evolution and biogeography of

216

bees, it is probable to assume that the basal lineages of Anthidiini happened

much earlier, possibly arose during the early Cretaceous, a period when the

continental connection of Africa and South-eastern Asia still existed. Further,

it is however possible that, more diversification within the tribe took place

much later (Eocene-Oligocene period), especially of genus Anthidium,

accounting for some of the fossils recorded in Engel and Perkovsky (2006).

A second and independent migration to the Neotropical region by the

probable ancestors of current Neotropic genera is conceivable. Such an event

probable took place when the gap between the drifted continents of South

America and Africa was narrow enough to permit short insect flight along its

corridors and similar climatic conditions still existed that provides similar

forage resources for these bees.

Evolution of parasitism within the tribe Anthidiini probably followed

this scenario. The parasitic genera probably evolved ones within the lineages

of the Anthidiini. This lineage comprises mainly two Afrotropical and Oriental

genera (Afrostelis and Euaspis), and a Neotropic genus Hoplostelis.

Revision of the Afrotropical Genus Anthidiellum

In the course of this present study, examination of all the Afrotropical

species of genus Anthidiellum (except A. madli Pauly, A. mediale Pasteels, A.

micheneri Pauly and A. transversale Pasteels) revealed two undescibed species

Anthidiellum (Pycnanthidium) frontoreticulatum spec nov and Anthidiellum

(Pcynanthidium) krugerense spec nov. A. frontoreticulatum spec nov, was

217

collected by Michener in 1967 from Kenya. However, he did not describe and

published this species as spec. nov, hence the present study has provided

description on this new species. A. krugerense spec nov, was found among the

bee collections of the South Africa National Collection. The only specimen

found was collected from Kruger National Park, South Africa, hence this new

species has been named after the place of collection. A. krugerense differ from

other Anthidiellum species by having lateral margin of scutum (in dorsal view)

extended longitudinally into small carinae, with a groove separating the

extended surface with a larger carina next to the tegula.

During the revision of this genus, Anthidiellum auriscopatum was

found to differ significantly from Anthidiellum orichalciscopatum (males of

both species unknown) in the following characters. These include: A)

Extensive bright yellow maculation on face, legs and metasoma of A.

auriscopatum than in A. orichalciscopatum Antero-distal region of pronotum

gently rounded and non-lamellate, than strongly angulate and lamellate

pronotal lobe as in A. orichalciscopatum. C) Lower clypeal margin of A.

auriscopatum with six small papillae (projections), four projections on lower

clypeal margin of A. orichalscopatum. D) Hind basitarsus much short and

broader than hind tibia in A. auriscopatum, however in A. orichalciscopatum,

hind barsitarsus is long and nearly as broad as hind tibia. Based on the

variation of these characters between these species, I considered A.

auriscopatum and A. orichalciscopatum as two different species within the

subgenus Pycnanthidium Krombein.

218

In addition, Anthidiellum kimberleyana and A. spilotum which were

considered synonym by Pasteels (1984), have been re-stated into separate taxa

based on differences existing in the following characters: A) Reduced

integumental maculations on T3-T6 of A. kimberleyana than in A. spilotum. B)

Legs entirely brownish-black, lacking integumental maculations in A.

kimberleyana, which occurs in A. spilotum. C) Inner carina of mandible

distinct well developed in A. kimberleyana. D) Outer teeth of mandible appear

blunt in A. kimberleyana than in A. spilotum. E) Apical region of hind tibia

much broader in A. kimberleyana than in A. spilotum. During the revision of

the Afrotropical genus, the cladogram revealed two major clades of trees

within the genus. This apparently is not compatible with the subgeneric

classification of the genus (Michener, 2000). For the purpose of this study, the

subgeneric classification of Michener (2000) has been retained. The

Afrotropical species of genus Anthidiellum is postulated to be 17, excluding

the four species that were not available during the course of this study. A

taxonomic key to the Afrotropical species of the genus Anthidiellum has been

provided for the 17 species examined during this study.

219

CHAPTER SIX

CONCLUSIONS AND RECOMMENDATIONS

Conclusions and Recommendations

Of great interest to this current research are the possible contributions

of postulated re-classification and phylogeny of the tribe Anthidiini, as well as

providing a review of the Afrotropical species of genus Anthidiellum.

The monophyly of the Anthidiini and most of its genera were

confirmed. Further to this, three subtribes: Afranthidina, Anthidina and

Euaspina have been erected for the world’s anthidine genera. These comprise

the following: Subtribe Afranthidina—Afranthidium, Anthidioma, Aspidosmia,

Neanthidium; Subtribe Anthidina—Acedanthidium, Anthidiellum, Anthidium,

Bathanthidium, Benanthis, Dianthidium, Gnathanthidium, Hypanthidium,

Icteranthidium, Indanthidium, Notanthidium, Pachyanthidium,

Pleisianthidium, Pseudoanthidium, Trachusa; Subtribe Euaspina—

Anthidioctes, Afrostelis, Apianthidium, Aztecanthidium, Cyphanthidium,

Duckeanthidium, Eoanthidium, Epanthidium, Euaspis, Hoplostelis, Serapista.

Based on the result of the cladistic analysis and the biogeographic data

of the Anthidiini, an evolutionary scenario was postulated in which the

Anthidiini probably evolved in the early Cretaceous (over 100 million years

before present), with high diversification within the tribe occurring in the

Eocene era.

220

The Afrotropical and Oriental regions exhibit high bee richness within

the Anthidiini at the generic level, with 16 and 13 genera, respectively. One

subtribe, Afranthidina is limited to the African continent. High bee richness at

the generic level and basal groups of primitive forms are enough evidence to

postulate that, the origin and center of radiation for the Anthidiini probably

occurred in southern Pantropic Region (Southern Africa to South East Asia).

A second and independent migration to the Neotropical region by the

probable ancestors of current Neotropic genera of the Anthidiini is

conceivable.

The parasitic genera probably evolved ones within the lineages of the

Anthidiini. This lineage comprises mainly two Afrotropical and Oriental

genera (Afrostelis and Euaspis), and a Neotropic genus Hoplostelis.

Further, two new species Anthidiellum (Pcynanthidium)

frontoreticulatum sp nov and Anthidiellum (Pcynanthidium) krugerense sp

nov. have been described. Two synonyms A. auriscopatum = A.

orichalciscopatum and Anthidiellum kimberleyana = A. spilotum were re-

instated to their species status as separate taxa. A total of 17 species of

Anthidiellum were found to occur in Afrotropical Region and a taxonomic key

to these species has been provided as well.

Some gaps were identified during the course of this study that

affected analysis of the results, and these centred basically on the limited

221

number of specimens available for the analysis. It is therefore recommended

that

1. A global survey should be conducted for the world’s anthidine bees, in

order to ascertain all existing fauna, as well as increasing the numbers

of museum collections.

2. Phylogenetic studies of the Anthidiini, involving all known genera,

subgenera and possibly species, need to be conducted to accurately

confirm postulated theories in this present study.

3. Phenetic studies on these groups of closely related bees could be

conducted to establish their relationship at the molecular level.

222

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251

APPENDICIES

APPENDIX 1

Genera of Anthidiini in Each Series (** indicate parasitic groups)

Series A Series B

Acedanthidium Euaspis ** Afranthidium

Afrostelis ** Hoplostelis ** Anthidioma

Anthidiellum Hypanthidiodes Anthidium

Anthodioctes Hypanthidium Gnathanthidium

Apianthidium Icteranthidium Indanthidium

Aspidosmia Larinostelis** Neanthidium

Atropium Notanthidium Pseudoanthidium

Aztecanthidium Pachyanthidium Serapista

Bathanthidium Paranthidium

Benanthis Pleisanthidium

Cyphanthidium Rhodanthidium

Dianthidium Stelis

Duckeanthidium Trachusa

Eoanthidium Trachusoides

Epanthidium

(Michener, 2000)

252

APPENDIX 2

This give detailed information of the locality labels on each specimen examined

in this research. They have been captioned as material examined.

MATERIAL EXAMINED

Genus Acedanthidium Michener and Griswold

Type material: Acanthidium batrae Michener and Griswold, male and

female paratype: India, Uttar Pradesh, S.W.T. Batra (SEMC).

Genus Afranthidium Michener

Total specimens: 66 females and 45 males.

Afranthidium abdominale (Friese)

4 females & 2 males. SOUTH AFRICA: Pretoria, ii.1949, J.H. Stiglingh (1

female- SANC); Christiana, 9.i.1965, H.N. Empey (1 female- SANC);

Middlesburg, Kaoo, 11.ii.1971, H.N. Empey (1 female, 1 male- SANC); Tussen

Die Rivere Residence near Bethulie, 30º30'S 26º12'E, 30.iii.-3.iv.1987, C.D.

Eardley (1 female,1 male- SANC).

Afranthidium braunsi (Friese)

36 females & 16 males. BOTSWANA: Serowe, 4.xi.1984, P. Forchhammer (1

male- SANC); Serowe, 4.ix.1986, P. Forchhammer (1 female, 1 male- SANC);

Serowe, 1988, P. Forchhammer (2 males- SANC). NAMIBIA: Near Orange

River, 14 Km S Rosh Pinash, 28º17'S 16º55'E, 27.x.1974, R.H. Watmough (1

female- SANC). SOUTH AFRICA: Beaufort West, Cape Town, 24.xi.1933,

J.Ogilvie (1 female- SANC); Mondeor, Johannesburg, 4.xii.1961, H.N. Empey

(1 female- SANC); Wepener, 18.i.1965, H.N. Empey (1 female- SANC);

Middleburg, 16.xi.1972, 24.i.1973, E. Holm (5 females- SANC); Oudtshoorn

253

Experiment Farm, 4.iii.1981, R.H. Watmough (3 female- SANC); Petrsville,

30º05'S 24º14'E, 2.xi.1981, H. Horn (1 female- SANC); Pretoria District,

Soutpan, 25º24'S 28º06'E, 13.xii.1982, C.D. Eardley (1 female, 3 males-

SANC); Rooiheuwel, 5 Km E Oudtshoorn, 33º36'S 22º17'E,5.x.1981, xi.1982,

2.xii.1983, 2.xii.1984, R.H. Watmough (8 females, 5 males- SANC);

Magaliesberg, Castle Gorge, 25º48'S 27º34'E, 17.ii.1993, R. Urban (1 male-

SANC); Adullam Farm, 28º32'S 28º28'E, 15-18.i.1986, 17-18.i.1994, C.D.

Eardley (4 females- SANC); 20 km E Waterpoot, 22º52'S 29º47'E, 6.i.2004,

C.D. Eardley (1 male- SANC); Guateng, Soutpan, 25º24'S 28º06'E, C.D.

Eardley (1 male- SANC).

Afranthidium folliculosum (du Buysson)

7 females and 14 males. KENYA: Central Island, Lake Turkana, 5.vii.1980, 23-

25.vii.1980 (12 males- KNM). NAMIBIA: Okahandija District, Erichsfelde 44,

21º35'S 16º56'E, 20-22.iii.2003, A.H. Kirk-Spriggs (1 female- SMWN).

SOUTH AFRICA: Alberton, Transvaal, 28.i.1966, H.N. Empey (1 male-

SANC); D’ Nyala Nature Reserve, Ellisras District, 23º45'S 27º49'E, 10-

14.xi.1986, V.M. Uys (1 female- SANC); D’ Nyala Nature Reserve, Ellisras

District, 23º45'S 27º49'E, 8-12.xii.1989, C.D. Eardley (3 females, 1 male-

SANC); Hotazel, 27º13'S 22º57'E, C.D. Eardley (1 female- SANC).

Afranthidium immaculatum (Smith)

7 females and 6 males. KENYA: Makalia Bridge Area, Lake Narkura,

28.xi.1977, E. Mlen (1 female- KNM); Gilgil, Aloe-Lalashwa grassland,

9.v.1982 (1 female, 1 male- KNM). SOUTH AFRICA: Graskop, E. Transvaal,

6.v.1964, H.N. Empey (1 male- SANC); Orange Free State, Marquad,

26.xii.1975, H.N. Empey (1 male- SANC); Long Tom Pass, 25º07'S 30º35'E,

254

i.1977, E.F.Whiteside (1 female- SANC); Tussen Die Rivere Residence near

Bethulie, 30º30'S 26º12'E, 30.iii.-3.iv.1987, 20-26.xii.1994, C.D. Eardley (2

females- SANC); Lesotho, 12km NE of Mamohau, 23.i.1992, H. Geertsema (1

female-SANC); Tussen Die Rivere Residence near Bethulie, 30º30'S 26º12'E,

21.i.1994, R. Adam (1 male- SANC). ZIMBABWE: Chimanimani Mountain

Hut, 4.iv.1972, R.H. Watmough (1 female- SANC).Afranthidium minutulum

(Friese)

12 females and 7 males. NAMIBIA: Ugab Road, 2 km W Brandberg, 20º57'S

14º06'E, 22-24.x.1998, E. Marais and A. Kirk- Spriggs (1 female, 2 males-

SMWN). SOUTH AFRICA: Augrabies National Reserve, 11.xii.1974, H.N.

Empey (1 female, 1male- SANC); Winheok, 22º34'S 17º06'E, xii.1983,

28.xiii.1983, R. Oberprieler (7 females, 4 males- SANC); 20 km E of

Waterpoort, 22º52'S 29º47'E, 6.i.2004, C.D. Eardley 2 females- SANC);

Richsterveld National Park, near Oena Mines, 28º03'S 17º03'E, R. Combey (1

female-SANC).

Genus Afrostelis Cockerell

Total specimens examined: 21 females and 10 males.

Afrostelis aethiopica (Friese)

19 females and 7 males. DEMOCRATIC REPUBLIC OF CONGO:

Bulawayo, S. Rhodesia, xii.1912, G. Arnold (1 female- TMSA). SOUTH

AFRICA: Lady Grey, 13.xii.1924 (1 male- NHML); Katberg, 15-30.i.1933,

R.E. Turner (1 female- NHML); Discovery Florida Transvaal, 16.ix.1958, H.N.

Empey (1 male- SANC); Mogoto National Reserve, Zebediela, 24º15'S

29º13'E, 22-25.x.1979, C.D. Eardley (1 male- SANC); Groenfontein, 35 km E

255

Thabazimbi, 24º34'S 27º45'E, 27.xi.1980, C. Kok (1 male- SANC);

Rustenburg Nature Reserve, 25º40'S 27º12'E, 17-20.iii.1980, C.D. Eardley (1

female, 1 male- SANC); Duiweiskloof, 23º42'S 30º06'E, 12-14.i.1987, C.D.

Eardley (1 male- SANC); Molteno Pass, near Beaufort West, 32º12'S 22º33'E,

14.xii.1988, C.D. Eardley (1 female- SANC); Karoo National Park, Beaufort

West, 32º20'S 22º30'E, 13.ii.1991, C.D. Eardley (1 male- SANC); Adullam

Farm, 28º32'S 28º28'E, 15-18.i.1986, 17-18.i.1994, C.D. Eardley (4 females, 1

male-SANC); Adullam Farm, 28º32'S 28º28'E, 18.i.1994, V.M.Uys (1 female-

SANC).

Afrostelis tegularis (Cockerell)

Type material: Afrostelis tegularis (Cockerell), female holotype:

Kamaiola, Congo, MRAC.

Additional material: CONGO: Bukama, 6.v.1911, Dr Bequaert (1 male-

MRAC); Luluabourg, 12.v.1919, P.Callewaert (1 male- MRAC).


Total material examined: 91 females & 68 males

Type material: Anthidiellum eritrinum Friese, female holotype, Gheleb

Eritrea, J. Pasteels 1963’, (HUMB).

A rubellum

1 female and 1 male. SERBIAN: (1female- HUMB). TUNISIA:

Tunis, .1976 (1 male- HUMB ‘labelled as type’).

A. flavescens

256

23 females and 8 males. ANGOLA: Bruco, 26.ii.-2.iii.1972 (1 male-

NHML). DEMOCRATIC REPUBLIC OF CONGO: Lonely Mine, S.

Rhodesia, 26.ii.1913, 21-24.v.1913, 13.xii.1913, 18.ii.1914, 26.xii.1914,

15.ii.1915, A.H. Swali (14 females- NHML). NAMIBIA: Kaoko Otavi, (1

female, 1 male- MRAC). ZIMBABWE: Bulawayo, 21.xii.1919, 28.xii.1924 (1

female, 2 males- SAMC, TMSA); Turk Mine, 16.xii.1957, H. Empey (5

females, 1 male- SANC); Premier Mine, 14.xii.1941 (1 female, 1 male-

SAMC).

A. absonulum

16 females & 17 males. BOTSWANA: Serowe, 17.ii.1984, 1988, P.

Forchhammer (2 females- SANC). DEMOCRATIC REPUBLIC OF CONGO:

Elizabethville, 11-17.ix.1931, J. Ogilvie (2 females- AMNH, MRAC),

3.xi.1933, M. Bequaert (1 male- MRAC); Haut-Uele: Paulis Mholi, iv.1947,

P.L.G. Benoit (1 female- MRAC). NAMIBIA: Salambala Forest, 17º09'57'S

20º52' 55'E, 26-27.xii.1998, A. Kirk-Spriggs & E. Marais, (1 female-

SMWN); idem, 23-29.xii.2002, A.H. & M.K. Kirk-Spriggs (7 females, 13

males- SMWN). SOUTH AFRICA: Mogol Nature Reserve, Ellisras District,

23º58'S 27º45'E, 25-26.i.1982, C.D. Eardley (1 female- SANC); idem,

21.xii.1987, B. Grobbelaar (1 female- SANC); D’ Nyala Nature Reserve,

Ellisras District, 23º45'S 27º49'E, 8.xii.1989, C.D. Eardley (1 male- SANC);

Schoongelegen, 24º12'S 27º45'E, 14.i.1991, V.M.Uys (2 males- SANC).

257

A. apicatum

7 females. SOUTH AFRICA: Happy Rest Nature Reserve, 22º55'57'S

29º44'06'E, 8.i.2004, C. D. Eardley (4 females- SANC). ZIMBABWE: Cashel,

30.xii.1947 (3 females- MRAC, SAMC).

Type material: Anthidiellum auriscopatum Strand, female holotype:

Spanish Guinea, Hinterland-Makomo, S.G. Tessman (HUMB).

Additional material: 2 females. DEMOCRATIC REPUBLIC OF THE

CONGO: Elizabethville, Piége Harris, 8.ix.1909, 11-17; idem, ix.1934, P.

Quarré (2 females- MRAC).

A. bipectinatum

5 females & 1 male: BOTSWANA: Serowe, 17.ii.1988, P.

Forchhammer (1 female- SANC). SOUTH AFRICA: Roodeplaat, xi.1978,

Pretoria District, C.D. Eardley (1 female- SANC); Soutpan, 25º24'S 28º06'E,

16.xi.1983, C.D. Eardley (2 females- SANC); Rustenburg Nature Reserve,

25º40'S 27º12'E, 8.xii.1983, C. D. Eardley (1 male- SANC).

A. bulaweyense

2 males. DEMOCRATIC REPUBLIC OF CONGO: Uamgebiet,

Bosum, 21.iii.1931, S.G. Tessmann (1 male- HUMB). ZIMBABWE:

Bulawayo, xii.1938 (1 male- SAMC).

Type material: Anthidiellum frontoreticulatum (sp. nov), holotype

female: Kenya, 26.v.1967, C.D. Michener, MRAC; Anthidiellum

frontoreticulatum (sp. nov), allotype male: Kenya, 26.v.1967, C.D. Michener,

MRAC; idem paratypes 2 females, 1 male, SEMC.

258

Type material: Anthidiellum kimberleyana (Friese), holotype female:

Kimberley, South Africa, Bro. Power (HUMB).A. orichalciscopatum

1 female. SOUTH AFRICA: Mossel Bay, i.1940 (1 female- SAMC).

A. otavicum

6 females and 3 males. KENYA: South Melindi, 26.v.1967, C. D.

Michener (4 female, 2 males- MRAC, AMNH). NAMIBIA: Otavi,

27.xii.1933, 16.i.1934, J. Ogilvie (2 females, 1 male- NHML).

Type material: Anthidiellum (Pycnanthidium) polyochrum

Mavromoustakis, holotype female: Kaross, South Africa, SAMC.

Additional material: 1 female, 1 male: NAMIBIA: 40 km North

Usakos, 18.ii.1977, J. G. & B. L. Rozen (1 male- AMNH); Ameib Farm,

19mls, NW Karibib, 31.i.-2.ii.1972 (1 female- NHML).

A. spilotum

5 females and 7 males: ANGOLA: Bruco, 26.ii.-2.iii.1972 (1 female-

NHML). BOTSWANA: Serowe, 17.ii.1988, P. Forchhammer (1 female-

SMWN). SOUTH AFRICA: Mossel Bay, i.1940 (1 female- SAMC); Crystal

Waters, Breedsnek, 4.xi.1962, H.N. Empey (2 females, 5 males- SANC);

Queenstown, 16.xii.1979 H.N. Empey (1 female, 1 male- SANC);

Saartjiesnek, 25º41'S 27º54'E, 12.xi.1983, C. Bellamy (1 male- SANC).

Type material: Anthidiellum (Pycnanthidiellum) sternale Pasteels,

holotype male and 1 paratype male: Breedsnek Transvaal, South Africa,

259

TMSA; Anthidiellum (Pycnanthidiellum) sternale Pasteels, 1 paratype male:

Mogal Nature Reserve, South Africa SANC).

Additional material: 1 female & 1 male. SOUTH AFRICA: Nylsvley

Nature Reserve, 24º39'S 28º42'E, 10-11.xii.1979, C.D. Eardley (1 male-

SANC); Mogal Nature Reserve, Ellisras District, 23º58'S 27º45'E, 25-

26.i.1982, C.D. Eardley (1 female- SANC).

A. tegwaniense

2 female and 2 males. ZIMBABWE: Bulawayo, 11.iv.1923, R.

Stevenson (1 male- TMSA); Khami, 28.x.1928 (1 female- SAMC); Khami,

27.x.1938 (1 female, 1 male- SAMC).

A. zebra

17 females and 21 males. BOTSWANA: Serowe, 17.ii.1984, 1988,

Forchhammer (1 male- SANC). SOUTH AFRICA: Willomore, 5.i.1905, H.

Brauns (2 female, 2 male- HUMB); Ben Alberts Nature Reserve, 24º37'S

27º23'E, 24-28.xi.1980, C.D. Eardley (1 male- SANC); Mogal Nature

Reserve, Ellisras District, 23º58'S 27º45'E, 25-26.i.1982, 27-29.ii.1984, C.D.

Eardley (1 female, 3 males- SANC); D’ Nyala Nature Reserve, Ellisras

District, 23º45'S 27º49'E, 19.xii.1987, 8.xii.1989, M. W. Mansell (6 females, 5

males- SANC), idem, 10-14.xi.1986, B. Grobbelaar (1 male); Breedsnek,

Crystal Waters, 17.xi.1962, (1 female, 3 males- SANC), H.N. Empey (1

female, 2 males- SANC), Pretoria, Soutpan, 25º24'S 28º06'E, 6.xi.1983, C. D.

Eardley (3 males- SANC); Happy Rest Nature Reserve, 22º55'57'S 29º44'06'E,

8.i.2004, C. D. Eardley (1 male- SANC).

260

Type material: Anthidiellum (Pcynanthidium) krugerense, holotype female-

SANC, South Africa, Kruger National Park, Skukuza, 24º59'S 31º35'E, 14-

17.i.1985, G.L. Prinsloo.

Genus Anthidioctes Moure

Anthidioctes willineri (Moure)

1 female and 1 male. BOLIVIA: Santa Cruz, Santiago, xi.1959 (SAMC).

Genus Anthidioma Pasteels

2 females. NAMIBIA: Diamond Area N º1, Pomona, 20.ix.1980, W.B.

Whitethead (SAMC).

Genus Anthidium Fabricus


Anthidium Cockerelli (Schwarz)

1 female and 1 male. UNITED STATE OF AMERICA: California, Nev Clark

Co, 3.3 rd mi SE Juanita Spr Ranch S of Riverside, 11-21.v.1983, F.D. Parker

and J.H. Parker (SANC).

Anthidium cordiforme (Friese)

9 females and 11 males. BOTSWANA: Farmers Brigade, 5 km SE of Serowe

Hiiside, i.1986, P. Forchhammer (1 males- SANC); Serowe, vi.1990,

Forchhammer (1 male- SANC). SOUTH AFRICA: Pretoria, 6-14.x.1971, E.

Holm (1 female- SANC); East Transvaal, Wolkberg, 30.viii.1975, R.H.

Watmough (1 female- SANC); Buysdor, 23º01'S 29º22'E, 16.i.1980, C.Kok (1

female- SANC); Wylliespoort, Ingwe Motel, 22º58S 29º57'E, 20-22.i.1982,

261

C.D. Eardley (1 female- SANC); Eureka City, Louw Creek SE , 3.iv.1983,

S.L. Chown (1 female- SANC); Rustenburg Nature Reserve, 25º40'S 27º12'E,

14.iii.1984, M. Mansell (1 female- SANC); 14-29 km W Alldays, 22º44'S

28º57'E, 14.iii.1990, C.D. Eardley (2 males- SANC)Farm Carpe Diem, near

Trichardtsdal, 24º10'S 30º26'E, 30-31.ix.1994, C.D. Eardley & M. Mansell (1

female- SANC); Faerie Glen, Pretoria, 25º44'S 28º13'E, 25.ii.1998, M. Smit (1

male- SANC); Gauteng, Soutpan, 25º24'S 28º06'E, 11.iii.1998, S. Muller (1

male- SANC); Northern Province, Thabahaswa, Groenkom Farm, near

Potgietersrus, 24º03'S 29º02'E, 21-23.ii.2001, M. Stiller (1 female- SANC);

Roodeplaat Research Station, 25º35'S 28º21'E, iii.2004, C.D. Eardley (1 male-

SANC); Gauteng, Soutpan, 25º24'S 28º06'E, 2.iii.2005, C.D. Eardley (1

female, 2 males- SANC); Happy Rest Nature Reserve, 22º55'S 29º44'E,

8.i.2004, C.D. Eardley (1 male- SANC).

Anthidium echinatum (Klug)

1 female and 1 male. MAURITANIA: Akjoujt, Biological Integrated Lucust

Control, xi.1993-ii.1994, J. Tabel (SANC).

Anthidium niveocinctum (Gerstaecker)

4 females and 3 males. SOUTH AFRICA: Waterpoort, SE 29º22'DC, iv.1978,

M. Botha (1 male-SANC); Groothoek Mine, 24º30 Ca, 6.iv.1980, A.P. du Toit

(1 female, 1 male- SANC); Langjan Nature Reserve, 23º52'S 29º14'E, 23-

24.i.1982, C.D. Eardley (1 female- SANC); Ranium, 25º05'S 28º17'E,

3.iii.1989, P. Snyman (1 female- SANC); Schuitdrift Road, 9 km SW

Tshipise, 22º40'S 30º04'E, 4.iii.1990, C.D. Eardley (1 female- SANC).

TANZANIA: Mkomazi Game Reserve, Ibaya, 03º58'S 37º48'E, 8.v.1995, A.

Russell (1 male- SANC).

262

Anthidium paroselae (Cockerell)

1 female and 1 male. UNITED STATE OF AMERICA: California, Nevada,

Clark Co, St Charles Gap, 16.v.1984, F.D. Parker (SANC).

Anthidium pontis (Cockerell)

3 females and 1 male. SOUTH AFRICA; Wilsonfontein, 22º40'S 15º40'E,

8.vii.1975, R.H. WAtmough (1 female-SANC); Langjan Nature Reserve,

23º52'S 29º14'E, 2.ii.1984, C.D. Eardley (1 female- SANC); Rus-en-Vrede,

near Oudtshoorn, 23º24'S 22º23'E, 18.xii.1986, M.W. Mansell (1 female-

SANC); Namaqualand, Hester Malan, 5.xi.1987, M. Struck (1 male-SANC).

Anthidium rodecki (Schwarz)

1 female and 1 male. UNITED STATES OF AMERICA: California, Nevada,

Humbodt Co, 10 mi N Winnemucca, 19.vi.1985, F.D. Parker (SANC).

Anthidium severini (Vachal)

13 females and 13 males. MALAWI: Senga Bay, 20 km NE of Salima, 7-

8.iii.1987, J. & A. Londt Woodland. SOUTH AFRICA: Mondeor,

Johannesburg, 21.x.1961, H.N. Empey (1 female- SANC); Retiefs Kloof,

Rustenberg, 13.iv.1963, H.N. Empey (1 female- SANC); Discovery Florida,

23.xii.1966, H.N. Empey (1 male- SANC); Kruger National Park, Tshokwana,

26.v.1969, L.C. Starke (1 female- SANC); Strubens Valley, florida, 15.xi.1970,

H.N. Empey (1 male- SANC); Pretoria, 9-17.xi.1971, E. Holm (1 male-SANC);

Nylsvley Reserve, x.1978, C.D. Eardley (1 female- SANC); Hennops River, 20

km W Pretoria, 25º74'S 27º55'E, 5.iii.1980, C.D. Eardley (1 male- SANC);

Groenfontein, 35 km E Thabazimbi, 24º34'S 27º45'E, 27.xi.1980, C. Kok (1

female, 1 male- SANC); Diamant, near Bulge Riviere, 24º07'S 27º33'E,

18.i.1983, M.W. Mansell (1 female, 2 males- SANC); Near Vivo, 22º56'S

263

29º33'E, 3.iii.1984, M.W. Mansell (1 male- SANC); Blyderiverspoortdam

Nature Reserve, 24º32'S 30º47'E, 25-26.x.1984, C.D. Eardley (1 female-

SANC); O.T.K Reserve, near Loskopdam, 25º27'S 29º24'E, 9.xii.1985, C.D.

Eardley (1 male- SANC); 5 km E of Thaba Nchu, 29º26'AB, 25.iv.1986, B.

Struckenberg & J. Londt (1 male- SANC); Malta Forest near The Downs,

24º10'S 30º14'E, 3-6.iii.1986, C.D. Eardley (1 male- SANC); Mac Mac Falls,

10 km N Sabie, 25º02'S 30º48'E, 27.ii.1986, B. Grobbelaar (1 female- SANC);

Swadini, Blydepoort Nature Reserve, 24º32'S 30º54'E, 26-29.i.1987, B.

Grobbelaar (1 male- SANC); Langjan Nature Reserve, 23º52'S 29º14'E,

10.iii.1990, C.D. Eardley (1 male- SANC); 14-29 km W Alldays, 22º44'S

28º57'E, 14.iii.1990, C.D. Eardley (4 female- SANC).

Genus Apianthidium Pasteels

1 male. MALAYSIA: Borneo, Sarawak, Mt Matang, 13.xii.1913, G.E.

Bryant (NHML).

Genus Aspidosmia (Brauns)

Type material: Aspidosmia arnoldi (Brauns), lectotype female and

male paralectotype. Willowmore, South Africa, TMSA.

Genus Aztecanthidium Michener and Ordway

Type material: Aztecanthidium tenochtitlanicum Snelling, male and

female paratype: Mexico, Jalisco, C.D. Michener (SEMC).

264

Genus Bathanthidium Mavromoustakis

1 female. SIBERIA: Kudia River, Amagu, vii.1923 (NHML).

Genus Benanthis (Pasteels)

Benanthis madagascariensis (Benoist), 1 male, A. Sekrig, MRAC.

Genus Cyphanthidium Pasteels

Cyphanthidium sheppardii (Mavromoustakis).

1 male. DEMOCRATIC REPUBLIC OF CONGO: Bassin Lukuga,

1934, De Saeger (MRAC).

Genus Dianthidium (Cockerell)


Dianthidium curvatum (Cockerell)

1 female and 1 male. UNITED STATES OF AMERICA: Utah, Delta,

3.viii.1951, G. E. Bohart (1 female- SANC); Utah, Cacheco, 5 km W Paradise,

4.ix.1987, D. K. Broemeling (1 male, SANC).

Dianthidium cressonii (Dalla Torre)

1 female and 1 male. UNITED STATES OF AMERICA: Utah, Joab, Eureka,

10.viii.1965, G.L. Bohart (1 female- SANC); Utah, Farmington Cyn,

10.vi.1977, F.D. Parker (1 male- SANC).

Dianthidium dubium (Schwarz)

1 female and 1 male. UNITED STATES OF AMERICA: California, Shasta

CO, 2 km W Shingletown 2750´, 16-25.vii.1986, R. Miller (SANC).

265

Dianthidium pudicum (Cresson)

1 female and 1 male. UNITED STATES OF AMERICA: Utah, Farmington

Cyn, 30.vi.1970, P.F. Torchio (1 male- SANC); Utah, Uintah Co, SW

Bonanza, 5000'-5600 T10S R24E, 28.viii.1981, M. Schwartz & R. Miller (1

female- SANC).

Dianthidium subparvum (Swenk)

1 female and 1 male. UNITED STATES OF AMERICA: California, Lassen

Co, 45 m NW, Susanville, 23.vii.1990, P.F. Torchio, G. Trostle (SANC).

Dianthidium ulkei perterritum (Cockerell)

2 females and 2 males. UNITED STATES OF AMERICA: Arizona, Cochise

Co Cave Cr.., 5 km SW, Portal, 4.ix.1986, F. Parker (1 female, 1 male-

SANC); California, Shasta CO, 20 mi N Burney, 25.vii.1990, P.F.Torchio &

G. Trostle (1 female, 1 male- SANC).

Dianthidium parvum (Cresson)

1 female and 1 male. UNITED STATES OF AMERICA: Utah, Uintah Co,

7.ix.1965, R.W. Rust (SANC).

GENUS Duckeanthidium MOURE AND HURD

Type material: Duckeranthidium theieli Michener, male and female

paratype: Costa Rica, R. Thiele (SEMC).

GENUS Eoanthidium POPOV

Total number of specimens examined: 20 females and 13 males.

266

Eoanthidium armaticeps (Friese)

3 female and 2 males. DEMOCRATIC REPUBLIC OF CONGO:

Elizabethville, 31.v.1935, P. Quarré (1 female- MRAC); Elizabethville,

29.iii.1938, H.J. Brédo, (1 female- MRAC). ZIMBABWE: Springvale, S.

Rhodesia, 26.xii.1911 (1 male- SAMC); Lonely Mine, S. Rhodesia, 9.ii.1919

(1 female- SAMC); Sawills, S. Rhodesia, 12.x.1923 (1 male- SAMC).

Eoanthidium bituberculatum (Pasteels)

1 female and 5 males. NAMIBIA: Brandberg, Snusib Ravine, 21º04'S 14º36'E,

2.xii.2000, E. Marais & W. Mey (1 female- SMWN). SOUTH AFRICA:

Spritzkoppe, 17 km Usakos, 21º59'S 15º35'E, 5.vii.1975, 18.vii.1976, R.

Watmough (4 males- SANC); Richtersveld Paradys Kloof, 28º19'S 17º00'E,

11.v.1990, M. Stiller (1 male- SANC).

Eoanthidium rothschildi (Vachal)

14 females and 4 males. NAMIBIA: Etosha, Ondundo Hills, 19º15'S 15º43'E,

10.x-30.xi.1986, E. Griffin. SOUTH AFRICA: Mogal Nature Reserve, Ellisras

Dist, 23º58'S 27º45'E 27-29.ii.1984, C.D. Eardley (1 female- SANC);

Rustenburg Nature Reserve, 22º40'S 27º12'E, 23-26.ii.1981, C.D. Eardley (1

female- SANC); Presidentsrus, near Witbank, 25º41'S 29º22'E, 20.iii.1986,

V.M. Swain (1 female- SANC); Kruger National Park, Pafuri, 22º26'S 31º12'E,

9.iv.1986, L.E.O. Braack (1 female- SANC); 14-29 km W Alldays, 22º44'S

28º57'E, 14.iii.1990, C.D. Eardley (9 females, 3 males- SANC); Between

Nylstroom and Warmbaths, 24º43'S 28º18'E, 18.i.1991, M. Harney (1 female-

SANC).

267

Eoanthidium (Clistanthidium) turnericum (Mavromoustakis)

2 females and 2 males. SOUTH AFRICA: South West Africa, Zesfontein (1

female, 1 male- SAMC); South West Africa, Namib Park, 23º28'S 15º19'E,

16.iii.1983, C.D. Eardley (1 male- SANC); Richtersveld National Park,

28º18'S 16º58'E, 12-14.ix.2001, C.D. Eardley (1 female- SANC).

GENUS Epanthidium MOURE

Epanthidium trigrinum (Schrottky)

1 female and 1 male. ARGENTINA: Salta, El Allsal, i.1990, iv.1991, Fritz

(AMNH).

GENUS Euaspis GERSTAECKER

Total number of specimens examined: 35 females and 5 males.

Euaspis abdominalis (Fabricus)

34 females and 4 males. GHANA: University of Cape Coast, Cape Coast,

05º07'N 01º15'W, 13.ii.2003, N. Akrasi (2 females- SANC). SOUTH

AFRICA: Tongaat, viii.1909, H.C. Burnup (1 female, 1 male- SANC);

Knauzl, 21.xi.1916, Marley (1 female- SANC); Maritzburg, 12.i.1917, C.

Akorman (2 female-SANC); Pretoria, Transvaal, 4.xii.1919 (1 female-SANC);

Gever, Transvaal, 12.xii.1931 (1 female- SANC); Pietersburg, Pretoria,

xii.1961, H. Botha (1 female- SANC); Strubens Val. Transvaal, 5.xii.1961,

H.N. Empey (1 female- SANC); Stanger, Natal, 14.iii.1963, H.N. Empey (1

male- SANC); Nelspriut, xii.1962, iv.1964 (2 females-SANC); Mpumalanga,

Graskop, Mrt. 1969 (2 females- SANC); Pretoria, Retief, i.1970, S. Kohmeyes

(1 female- SANC); Pretoria, P.A. Battey, xii. 1972 (1 female- SANC); Mogoto

268

National Reserve, Zebediela, 24º15'S 29º13'E, 22-25.x.1979, C.D. Eardley (1

female- SANC); Mogoto National Reserve, Zebediela, 24º15'S 29º13'E, 12-

14.iii.1980, G.L. Prinsloo (1 female- SANC); Charters Creek, St Lucia,

28º12'S 32º25'E, 14-16.i.1981, M.W. Mansell (1 female- SANC); Fanies

Island Camp, 28º10'S 32º25'E, 14-16.i.1981, G.L. Prinsloo (1 male- SANC);

O.T.K. Reserve, near, Loskopdam, 25º27'S 29º24'E, 9-11.xii.1985, C.D.

Eardley (1 female- SANC); Cathedral Peak Area above Mike 1s Pass, 28º59'S

29º14'E, 28-29.iii.1985, C.D. Eardley (1 female- SANC); Ludwig, Pretoria,

6.xii.1986 (1 female- SANC); Kosi Bay, 26º58'S 32º48'E, 10-11.ii.1990, C.D.

Eardley (3 females, 1 male- SANC); Kosi Bay, 26º58'S 32º48'E, 10-

11.ii.1990, N. Verheijen (2 females- SANC); Happy Rest, 22º55'S 29º44'E,

8.i.2004, C.D. Eardley (1 female-SANC). ZIMBABWE: Bindura, Rhodesia,

1938 (1 female- SANC); Salisbury, Rhodesia, A. Watsham (1 female-

SAMC).

Euaspis erythros (Meunier)

1 female and 1 male. SOUTH AFRICA: Natal, 1.xii.1930 (1 male- SANC);

Barberton, Transvaal, xii.1978, C.D. Eardley (1 female- SANC).

GENUS Gnathanthidium PASTEELS

3 females. KENYA: Shimoni, iv.1955 (1 female- MRAC).

ZIMBABWE: Bulawayo, S. Rhodesia, 28.xii.1924 (1 female- MRAC);

Nyamandhlovu, S. Rhodesia, 9.i.1960 (1 female- MRAC).

269

GENUS Hoplostelis DOMINIQUE

1 female and 1 male. ARGENTINA: Salta, Cafayate, i. 1992, Fritz

(AMNH).

GENUS Hypathidiodes MOURE

Hypanthidioctes argentina (Urban).

1 female and 1 male. ARGENTINA: Salta, Rosario Lerma, xi. 1993, Fritz, (1

male- AMNH); Salta, El Allsal, iii.1992, Fritz (AMNH).

Hypanthidioctes emarginatum (Urban)

1 female and 1 male. UNITED STATE OF AMERICA: Utah, Uintah Co, SW

Bonanza, 5000'-5600 T10S R24E, 28.viii.1981, M. Schwartz & R. Miller

(SANC).

GENUS Hypanthidium COCKERELL

Hypanthidium flavomarginatum Smith

2 females and 2 males. ARGENTINA: Mendora, 20.x.1908 (TMSA).

Genus Icteranthidium Latreille

Icteranthidium laterale Latreille

1 female & 1 male. ITALY: Ulten, Tirol, Sclett (1 male-SEMC). RUSSIA:

Chotin-Slov, 4.viii.1950, C. Pàdr (1 female-SEMC).

Genus Indanthidium Michener And Griswold

1 female & 1 male. INDIA: Poona, Lonavia, 20.i.1959, F.L. Wain (SEMC)

270

Genus Neanthidium Pasteels

1 female and 1 male. LIBYA: Jebel Soda, Wadi Ghodaifa, 3iii.1952,

4.iii.1952, K.M. Guichard (NHML).

Genus Notanthidium (Isensee)

Notanthidium steloides (Spinola)

1 female and 1 male. CHILE: Canelo, Santiago, 31.i.1949, L.E. Pena (1 male-

SAMC); Ceigual, 1400-1640 m E, 13.i.1955, L.E. Pena (1 female- SAMC).

Genus Pachyanthidium Vachal

Total number of specimens examined: 14 females & 35 males.

Pachyanthidium ausense (Vachal)

3 females & 6 males. BOTSWANA: Farmers Brigade, 5 km SE of Serowe

Hiiside, 1988, xi.1989, P. Forchhammer (2 females- SANC). NAMIBIA: Near

Orange River, 14 Km S Rosh Pinash, 28º17'S 16º55'E, 27.x.1974, R.H.

Watmough (5 male- SANC). SOUTH AFRICA: Richterveld National Park,

Hellskloof Pass, 9.x.1974, H.D. Brown (1 female- SANC); Namaqualand

2917 DB, 26.ix.1987, N.R. Hester Malan, M. Struck (1 female- SANC).

Pachyanthidium benguelense (Vachal)

6 females & 12 males. GHANA: Kakum Conservation Area, 05º20'N

001º22'W, 17.ii.2005, R. Combey (1 female- UCCEM). DEMOCRATIC

REPUBLIC OF CONGO: Elizabethville, 20.ix.23, 8.iv.1933, M. Bequaert (2

males- MRAC); Umtali District, Witchwood Valley, 27.i.1959, A.C. van

Bruggen (1 female- SANC); Salisbury, A. Watsham (1 female- SANC).

MALAWI: Fort Lister, Mt Mulanje, 100m SE, 15.35 DC, 2-4-xii.1983 (1

271

female- SANC). SOUTH AFRICA: Silverton, 6.i.1962, H.N. Empey (1 male-

SANC); Swartkops, 19.x.1963, H.N. Empey (1 female- SANC); Natal,

Milands, Escourt, 5.ii.1975, H.N. Empey (1 male- SANC); Schoemanville,

Dam Wall, 12.x.1975, H.N. Empey (2 males- SANC); Drakenberg, Cathkin

Peak District, 26.x.1976, H.N. Empey (1 male- SANC); Nylsvley Reserve,

x.1978, C.D. Eardley (1 female- SANC); Mogol Nature Reserve, Ellisras

District, 23º58'S 27º45'E, 25-26.i.1982, C.D. Eardley (1 female, 1 male-

SANC); idem, 3.xii.1985, J.S. Donaldson (1 male- SANC); Eastern Transvaal

Cooperative Reserve, near Loskopdam, 25º27'S 29º24'E, 9-11.xii.1985 C.D.

Eardley (1 male- SANC); Guateng, Soutpan, 25º24'S 28º06'E, 2.iii.2005, C.D.


Pachyanthidium bicolor (Lepeletier)

1 female & 3 males. GHANA: Kakum Conservation Area, 05º20'N 001º22'W,

18.i.2005, R. Combey (1 male- UCCEM). UGANDA: Entebbe, 30.vii.1913,

C.C. Gowdey (1 male- SANC); 20 km W Kampala, v.1952, E. Pinbey (1

male- KNM); Bwamba, W. Uganda, ii-iii.1957, R. Carcasson (1 female-

KNM).

Pachyanthidium cordatum (Smith)

4 females & 15 males. DEMOCRATIC REPUBLIC OF CONGO: Salisbury,

A. Watsham (1 female- SANC). SOUTH AFRICA: Maritzburg, 23.ii.1917, C.

Akerman (1 female- SANC); Pretorius Kop, 27.x.1950, H.K.Munro (1 male-

SANC); Scottburgh, 15.iii.1963, H.N. Empey (1 female, 1 male- SANC);

Stijdom Tunnel, 19.ii.1969, L.C. Starke, (1 male- SANC); Natal, Tugela

Ferry, 13.xi.1971, H.N. Empey (1 male- SANC); Umtentwini, 20.iv.1973,

H.N.Empey (2 males- SANC); Natal, Hluhluwe Game Reserve, 27º54'S

272

32º22'E, 8.ii.1974, R.H. Watmough (1 male- SANC); Assegaaihoutkloof, near

Groblersdal, 25º15'S 29º25'E, 4.xi.1979, R.H. Watmough; Mogol Nature

Reserve, Ellisras District, 23º58'S 27º45'E, 19-23.xi.1979, S.J. van Tonder, C.

Kok, G.L. Prinsloo & M.W. Mansell (1 male- SANC); Leboea, Mahlasi, 18

km NE Burgersfort, 24º31'S 30º25'E, 24.xi.1981, G.L. Prinsloo (1 male-

SANC); Kuleni Farm, Hluhluwe, 27º54'S 32º22'E, 14.ii.1990, C.D. Eardley (1

male-SANC); Mfongesi, Zululand, W.E.Jones (2 males-SANC); Tang Terr,

Ukerewe, Father Conrads (1 male- SANC). UGANDA: 14 Falls, Thika,

xi.1959, R. Carcasson (2 males- KNM).

Genus Paranthidium Cockerell and Cockerell

1 female and 1 male. UNITED STATES OF AMERICA: Arizonia,

Onion Saddle, Chiricahua Mts., Cochise, 4.ix.1967 (1 male- SANC); Arizona,

Cochise Co., South Western Research Station, 5 mi W Portal, 24.viii.1973 (1

female- AMNH).

GENUS Pleisianthidium CAMERON


Plesianthidium bruneipes (Friese)

5 females & 2 males. SOUTH AFRICA: Namaqualand, Hester Malan,

20.viii.1985, 13.ix.1987, M. Struck (2 females, 1 male- SANC); Kamiesberg

Pass, 30º12'S 17º59'E, 11. ix.1987, C.D. Eardley (1 female, 1 male- SANC);

Farm Dassiefontein, near Kamieskroon, 30º09'S 17º59'E, 1.x.1990, C.D.

Eardley (2 females- SANC).

273

Plesianthidium calescens (Cockerell)

14 females & 21 males. SOUTH AFRICA: Clanwilliam District, Biedouw

Valley, 32º08'S 19º14'E, 5-7.ix.1987, C.D. Eardley (6 females, 5 males-

SANC); Namaqualand, Hester Malan, 20.viii.1985, 14.x.1987, M. Struck (2

females, 3 males- SANC); Pakhuis Pass, 32º08'S 19º02'E, 7.ix.1987, C.D.

Eardley (1 female, 5 males- SANC); Farm Dassiefontein, near Kamieskroon,

30º09'S 17º59'E, 1.x.1990, C.D. Eardley (5 females, 3 males- SANC);

Western Cape Province, near Skurfkop, 31º55'S 18º37'E, 20.ix.2005, C.D.


Plesianthidium cariniventre (Friese)

5 females & 16 males. SOUTH AFRICA: Pakhuis Pass, 32º08'S 19º02'E,

21.ix.1985, C.D. Eardley (1 female, 5 males- SANC); Namaqualand, Hester

Malan, 20.viii.1985, 2.ix.1986, 13.ix.1987, M. Struck (1 female,4 males-

SANC); Kamiesberg Pass, 30º12'S 17º59'E, 11. ix.1987, C.D. Eardley (1

female, 5 males- SANC); Nieuwoudtville Botanic Gardens, 31º22'S 19º07'E,

4.ix.1987, C.D. Eardley (1 male- SANC); Wuppertal, 32º17'S 19º13'E,

8.ix.1987, C.D.Eardley (1 male- SANC); Farm Dassiefontein, near

Kamieskroon, 30º09'S 17º59'E, 1.x.1990, C.D. Eardley (2 females- SANC);

Clanwilliam District, Biedouw Valley, 32º08'S 19º14'E, 27.ix.1990,

19.ix.2001, C.D. Eardley (1 female, 2 males- SANC).

Pleisianthidium fulvopilosum (Cameron)

3 females & 4 males. SOUTH AFRICA: Namaqualand, Hester Malan, 20-

28.viii.1985, 5.ix.1986, 12.viii.1986, 13.ix.1987, 16.ix.1987, 9.x.1987, M.

Struck (3 females, 4 males- SANC).

274

Pleisianthidium neli (Brauns)

5 females & 4 males. SOUTH AFRICA: 10 km W of Algeria, Clanwilliam

road, 32º21'S 19º03'E, 4.ix.1987, C.D. Eardley (1 female- SANC);

Kamiesberg Pass, 30º12'S 17º59'E, 11. ix.1987, C.D. Eardley (2 females, 3

males- SANC); Farm Dassiefontein, near Kamieskroon, 30º09'S 17º59'E,

1.x.1990, C.D. Eardley (1 female- SANC); Leliefontein, 30º23'S 18º16'E,

4.x.2003, C. Mayer (1 female, 1 male- SANC).

Pleisianthidium (Spinanthidium) trachusiforme (Friese)

28 females & 64 males. SOUTH AFRICA: Namaqualand, Hester Malan,

20.viii.1985, 2.ix.1986, 13.ix.1987, M. Struck (7 males- SANC); Clanwilliam

District, Biedouw Valley, 32º08'S 19º14'E, 5-7.ix.1987, C.D. Eardley (4

females- SANC); Kamiesberg Pass, 30º12'S 17º59'E, 11. ix.1987, C.D.

Eardley (1 male- SANC); O’okiep, 29º37'S 17º55'E, 13.ix.1987, C.D. Eardley

(2 males- SANC); Farm Dassiefontein, near Kamieskroon, 30º09'S 17º59'E,

1.x.1990, 16-17.ix.2001, C.D. Eardley (22 females, 46 males- SANC); Farm

Arkoep, 6 km N Kamieskroon, 30º19'S 17º56'E, 1-2.x.1990, C.D. Eardley (1

female, 3 males- SANC); 32 km N Nieuwoudtville, 31º16'S 19º46'E,

2.ix.2004, C.D. Eardley (1 female- SANC).

Plesianthidium (Spinanthidiellum) volkmanni (Friese)

27 females & 47 males. SOUTH AFRICA: Stellenbosch, 21.xii.1915, A.

Nicholson (4 females, 2 males- SANC); 5 km from Kuboos, 9.ix.1974, R.H.

Watmough (1 male- SANC); Gydo Pass, 33º14'S 19º20'E, 19.xi.1982,

26.ix.1990, C.D. Eardley (1 female, 4 males- SANC); Near Silverton, 33º34'S

18º23'E, 14.xi.1984, C.D. Eardley (1 male- SANC); Clanwilliam District,

Biedouw Valley, 32º08'S 19º14'E, 30.ix.1986, M.W. Mansell & J.H. Hoffman

275

(1 female, 1 male- SANC); Namaqualand, Hester Malan, 20.viii.1985,

2.ix.1986, 31.viii.1987, 13.ix.1987, M. Struck (7 females, 9 males- SANC);

Nieuwoudtville, 31º20'S 18º53'E, 6.ix.1986, H.C. Hatting (1 female- SANC);

Nieuwoudtville Botanic Gardens, 31º22'S 19º07'E, 9.ix.1987, C.D. Eardley (1

male- SANC); Wuppertal, 32º17'S 19º13'E, 8.ix.1987, C.D.Eardley (1 male-

SANC); Kamiesberg Pass, 30º12'S 17º59'E, 11. ix.1987, C.D. Eardley (1

male- SANC); 15 km E Citrusdal, 32º37'S 19º08'E, 4.x.1990, C.D. Eardley (1

female, 1 male- SANC); Farm Dassiefontein, near Kamieskroon, 30º09'S

17º59'E, 1.x.1990, 16-17.ix.2001, C.D. Eardley (3 females, 4 males- SANC);

Clanwilliam District, Biedouw Valley, 32º08'S 19º14'E, 22.ix.2001, C.D.

Eardley (1 male- SANC); Vanrhynsdorp, near Water Fall, , 31º47'S 18º45'E,

15.ix.2005, C.D. Eardley (5 males- SANC); Western Cape Province, near

Skurfkop, 31º55'S 18º37'E, 20.ix.2005, C.D. Eardley (3 males- SANC);

Western Cape, near Redeinghuys, 32º32'S 18º27'E, C.D. Eardley (1 female, 7

male- SANC); Die Berg, 32º32'S 18º27'E, 18.ix.2005, C.D. Eardley (3 males-

SANC).

Genus Pseudoanthidium Smith


Pseudoanthidium lanificum (Smith)

4 females & 2 males. DEMOCRATIC REPUBLIC OF CONGO: Bena Bendi,

v.1915, R. Mayné (1 female- MRAC); Abumombazi, 18-26.ii.1932, H.J.

Brédo (1 male- MRAC); Eala, 1932, A. Corbisier (1 female- MRAC);

Bambesa, 1933, H.J. Brédo (1 male- MRAC); Eala, xi.1935, viii.1938, J.

Ghesquiére (2 females- MRAC).

276

Pseudoanthidium tertium (Pasteels)

6 females & 4 males. KENYA: Nairobi, E. Shirley (1 female- KNM); Doho

Kosele, Kisii, xi.1950, Mrs Adamson (1 female- KNM); Makneni, v.1951,

B.T. Persons (1 female- KNM). SOUTH AFRICA: Katberg Mt, 32º31'S

26º38'E, 2.xii.1983, G.L. Prinsloo & N.C. Grobbelaar (1 male- SANC);

Lesotho, Katse, 24-28.xii.1991, H. Gertsema (1 female, 2 males- SANC);

Adullam Farm, 28º32'S 28º28'E, 17-18.i.1994, C.D. Eardley (1 female-

SANC); Kaffervoetpad, 15km S of Lydenburg, 25º14'S 30º29'E, 9-

10.xii.1995, R. Stals (1 female- SANC); Hlogoma, 29º48'S 29º32'E, 4-

7.xii.2001, S. van Noort (1 male- SANC).

Pseudoanthidium truncatum (Smith)

53 females & 25 males. DEMOCRATIC REPUBLIC OF CONGO: Eala,

28.ii.1939, J. Ghesquiére (1 female- MRAC); Ubangi, Nouvelle Anvers,

9.xii.1952, P. Basilewsky (1 female- MRAC); Tshuapa, Bokuma, vi, 1952,

R.P. Lootens (1 female- MRAC); Tshuapa, Bamania, ix.1954, R.P. Hulstaert

(1 female- MRAC). GABON: Gamba, Ogoué, 02º42'S 10º01'E, Syssou,

Ngoma, Moussavo (1 female, 1 male- SANC); GHANA: University of Cape

Coast, 05º07'N 01º25'W, N. Akrasi (3 females, 2 males- SANC); Cape Coast,

Antonio Gardens, 05º07'N 01º15'W, 3.xii.2003, C.D. Eardley (1 female, 1

male- SANC); Kakum Conservation Area, 05º20'N 001º23'W, 05º20'N

001º22'W, 18.i.2005, R. Combey (3 males- UCCEM). KENYA: Stoneham, E.

Surr Estate, 4.v.1943, 23.vi.1943, 15.vii.1948 (3 females, 1 male- KNM);

Amani, E. Usambara, iii.1950, E. Pinhey (1 male- KNM); Kakamega Forest

Reserve, v.1949, R.W. Barney (1 female- KNM); Kakamega Forest Reserve,

v.1950, Mrs. Adamson (1 female, 1 male- KNM); Yatta Kitui, iv.1960, R.H.

277

Carcasson (1 female- KNM); Karura, Nairobi, i.1972, M.P. Clifton (1 female-

KNM); Kakamega Forest Reserve, i.1980, M.P. Clifton (3 females- KNM);

Kakamega Forest Reserve, 00º24'N 34º50'E, v.2000, M. Kraemer & G. Velten

(1 male-KNM); Eastern Kerio Valley, 00º44'N 35º34'E, 3.vii.2003, D.

Martins, W. Kinuthia, C.D. Eardley & L. Njoroge (1 male-KNM); Nairobi

City Park, 31.vi.2006, M. Leah (1 male- KNM). SOUTH AFRICA: letaba

Estate, 3.viii.1947 (7 females- SANC); Water Val-Onder, 6.x.1962, H.N.

Empey (1 male- SANC); Malta Forest, 8.ix.1965, 13.ix.1965, 18.x.1974, H.N.

Empey (1 female, 1 male- SANC); Balloon Forest, 24º12'S 30º20'E, 8-

11.iv.1977, D.H. Jacobs (1 female- SANC); Scheeferskloof, near Loskop

Dam, 25º24'S 29º22'E, 3.vi.1978, r.h. Watmough (3 females- SANC); Kloof,

viii.1971, J. Yates (1 female- SANC); Nylsvley Reserve, x.1978, C.D Eardley

(2 females, 6 males- SANC); Barberton, xii, 1978, C.D. Eardley (1 female-

SANC); Pelgrimsrus, 24,30 DC, iv.1979, D.R. Swart (1 female- SANC);

Umzimkulu, 24.xii.1980 (1 female- SANC); De Kuilen, Lydenburg District,

25º10'S 30º32'E, 12.ii.1981, C. Moolman & W. Harrop (1 female- SANC);

Blyderiverspoortdam Nature Reserve, 24º32'S 30º47'E, 25-26.x.1984, C.D.

Eardley (1 female-SANC); Hoedspruit, 24º22'S 30º57'E, 23.xi.1985, S. King

(1 female- SANC); Duiwelskloof, 23º42'S 30º06'E, 15.xii.1985, J.S.

Donaldson (1 male- SANC); Entabeni Forest Reserve, 33º00'S 30º16'E, 7-

11.i.1987, C.D. Eardley (1 male- SANC); Ludwig, Pertorius, 17.i.1987 ( 1

female- SANC); Welgevonden, 24º52S 30º34'E, 11-20.xi.1990, V.M. Vys (1

female- SANC); Nelspruit, 30’25 BD, 11.iv.1994, J.D. Greeff (1 female-

SANC); Tembe Elephant Reserve, 27º03'S 32º25'E, 2.xi.1995. M. W. Mansell

(1 female- SANC); Roodeplaat, 25º41'S 28º18'E, 14.ii.1995, C.D. Eardley (1

278

female- SANC); Kiepersol, 26º10'S 31º07'E, 20.ii.1998, J. Smith (1 female-

SANC); Thabaphaswa, near Potgietersrus, 24º03'S 29º02'E, 21-23.ii.2001, M.

Stiller (1 male- SANC). UGANDA: Bwamba Forest, viii.1928, G.D.H.

Carpenter (1 female- KNM); Bwamba Forest, iii.1948, J.G. Williams (1

female- KNM).

Pseudoanthidium tuberculiferum (Friese)

15 females & 31 males. DEMOCRATIC REPUBLIC OF CONGO: 18 mi E of

Toues River, xii.1962 (1 female- MRAC). SOUTH AFRICA: Lesotho,

Teyateyaneng, 30.xii.1964 C. Jacot-Guillarmod (1 male-SANC); Cedarberg,

20.xii.1972, R.H. Watmough (1 female-SANC); Cathkin Peak, 26.x.1976,

H.N. Empey (1 male- SANC); Adullam Farm, 28º32'S 28º28'E, 20-26.ii.1980,

S.J. van Tonder (1 female- SANC); Natal, Underberg, 29º47'S 29º30'E,

17.xi.1981, S.J. van Tonder (1 male-SANC); Rooiheuwel, 5 km E

Oudtshoorn, 33º36'S 22º17'E, xi.1982, R. H. Watmough (1 male- SANC);

Georgida, 33º26'S 23º20'E, 25.xi.1983, G.L. Prinsloo & N.G. Grobbelaar (2

males- SANC); 20 km N of Silverton, 33º24'S 18º16'E, 14.xi.1984, C.D.

Eardley (4 males- SANC); Near Algeria Forest Reserve, 32º22'S 19º03'E,

14.xi.1984, C.D. Eardley (1 femlae, 1 male- SANC); Tweeling, 27º33'S

28º31'E, 15.i.1986, C.D. Eardley (1 male- SANC); Adullam Farm, 28º32'S

28º28'E, 15-18.i.1986, C.D. Eardley (3 females- SANC); Richtersveld

National Park, 5.x.1986, M. Struck (1 female- SANC); Verlatenkloof, near

Sutherland, 32º33'S 20º37'E, 10.xii.1988, C.D. Eardley (1 female, 2 males-

SANC); Mountain View, Karoo National Park, 32º15'S 22º32'E, 14.xii.1988,

C.D. Eardley (4 male- SANC); Gydo Pass, 33º14'S 19º20'E, 8.xii.1988, C.D.

Eardley (1 female, 1 male- SANC); Montagu Botanical Gardens, 33º47'S

279

20º09'E, 18.xii.1988, C.D. Eardley (1 female, 1 male- SANC); Doring River,

near Nieuwoudtville, 31º18'S 19º09'E, 17.xi.1984, 28.ix.1990, C.D. Eardley (1

female, 1 male- SANC); Arkoep Farm, 6 km N kamiekroon, 30º19'S 17º56'E,

1-2.x.1990, C.D. Eardley (1 female, 1 male- SANC); Pakhuispas, near

Clanwilliam, 32º08'S 18º58'E, 27.ix.1990, C.D. Eardley (1 female- SANC).

Genus Rhodanthidium Spinola

Rhodanthidium septemdentatum (Latrielle)

1 female and 1 male. FRANCE: Var: Plan, vii.1979, 19.vii.1986, F. Schwartz

(AMNH).

Rhodanthidium siculum (Spinola)

2 females and 1 male. MOROCCO: Tanger (1 male-TMSA); Mogador,

ii.1905, escalera. UNITED STATES OF AMERICA: Florida, Bartagena, J.

Cac (1 female- TMSA).

Rhodanthidium fedtschericoi

1 female and 1 male. KAZAKHSTAN: Karataugeb, bei Djulek, Belamurun,

Koshantschilkow (1 female- AMNH). SERBARIA: Achtsha-Kujma,

7.vii.1934, V. Popov (1 male- AMNH).

GENUS Serapista COCKERELL

12 females & 20 males. KENYA: Stoneham, Nerang, 13.v.1952 (1

female-KNM); Bred, Ngong Escarb, xi.1957, P.R.O. Balley (1 male- KNM).

SOUTH AFRICA: Umtata, 15.xi.1920 (1 female, 7 males- SANC); Eshowe,

vi.1925 (1 female, 3 males- SANC); Pretoria, i.1944, Mr. Mogg (1 female-

SANC); Pertoria, i.1964, M.E.du T (1 male- SANC); 8 km N Nieuwodtville,

280

28.ix.1972, H.D. Brown, E. Koster & A. Prinsloo (1 female- SANC); Entabeni

Forest Reserve, 33º00'S 30º16'E, 7-11.i.1987, C.D. Eardley (1 female-

SANC);Groblershoop, 28º53'S 21º59'E, C. Moolman (1 female- SANC);

Langjan, SE 22’ 29 CC, 27.ii.1988, K. Kappmeier (1 male- SANC); Summit

of Mt Gilboa, KwaZulu-Natal, ii.2000, S.D. Johnson (1 female- SANC); 10

km N Concordia, 29º31'S 17º57'E, 23-24.viii.2006, R. Combey (2 females, 7

males- SANC). ZIMBABWE: Rhodesia, Chisawasha, 11.x.1979, A. Watsham

(1 male- SANC); Salisbury, A. Watsham (1 female- SANC).

GENUS Trachusa PANZER

Total number of specimens: 4 females and 7 males.

Trachusa aquilifilum (Strand)

3 females and 1 male.NAMIBIA: South West Africa, Otjimbumbe, iii.1923,

R. Kunene (2 females-SAMC, 1 male- MRAC); South West Africa, Ngosib

Koppie, Etosha National Park, 19º10'S 16º40'E, 9.iv.1989, E. Marais and J.

Irish (1 female- SAMC).

Trachusa schutedeni (Vachal)

2 males. CONGO: Costermansville, 1939, Dr. Hautmann (MRAC)

Trachusa serratula (Panzer)

1 female and 4 males. CHILE: Rostoch, Meche, 4.vi.1968, 28.v.1988,

20.vii.1988, Dr Brauns (TMSA).

Documents

Phylogenetic analyses of tribe Anthidiini