Taxonomic Studies of Collected in Khartoum State

By: Awatif Khidir Omer Yousif El Hag

A Thesis Submitted to the Department of Zoology infulfillment Master of Science in Zoology

Zoology Department, Faculty of Science University of Khartoum December, 200

© Awatif Khidir Omer

Dedication

To soul of my brother To my Beloved

Awatif

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Acknowledgements

It gives me a great pleasure to thank all those who contributed advice and help in this work. It is appropriate to mention several people who made suggestions, and provided editing. Deep gratitude is due to my supervisor Dr. Tigani M.H.Allam for his help, discussions and encouragement, thanks go to Dr. Sumaia Abukashwa for her help and advice. I would like to express my gratitude to Al-Zubair Mohammed Salih Prize for Innovation and Scientific Excellence for generous financial support. I am greatly indebted to Marcus Stuben (University of Wurzburg Germany, for providing me with some books and articles which were not available in . Many thanks to Dr. Mustafa Rizig from for his kind assistance, provision of books, and journals, for his keen guidance encouragement and his promise to help to put in the name of Sudan at the website of ants, thanks go to Dr. Collingwood of Liverpool Museum who identified some of the ants to species level. Of the all, thanks are especially extending to Dr. Brian Taylor (University of Nottingham- UK) for his great assistance in identification some of the ants to species level.

II Also thanks to Dr. Brian Fisher (California Academy of Sciences-USA), who identified some ants to the species level and imaged the ants to put it at the website www.antweb.org as ants of Sudan. I thank also Dr. Donat Agosti (American Musem of Natural History and Smithsonian Institution) who sent me the satellite map of Khartoum State, and thanks to his wife Livia Agosti (Deputy Head of the Political Affairs Division II African/Middle East) for delivering my specimens to Dr. Brain Fisher. Thanks to the Remote Sensing Centre (University of Khartoum) for locating collection site of the specimen in satellite map. Thanks to Dr. Abdullah Gomaa who gave me the GPS (Global Position System), to Dr. Zuhair Nour Eldeim for his advice and to Dr. Omran Fadl for his help. Not least but last, thanks are due to my colleagues in the Sudan Institute for Natural Sciences (SIFNS) for their encouragement and help with typing the thesis.

III Table of Contents Contents Page

Dedication……………………………………………………..……. I Acknowledgements………………………..…………………….….. II Table of Contents……………………………………………………. IV List of Tables…………………………………………..………...... IX List of Figures……………………………………..…………….….. X List of Plates………………………………………………………… XI list of Appendices XIII ...... Abstract………………………………………….…………….……. XIV Chapter 1: Introduction………………...…………………….… 1

1.1. Social Organization…………………….………………..…….. 1

1.2. Feeding ………………………………………………...…. 2 1.3. Objective of this study ……………………………………..….. 3 Chapter 2: Literature Review…………………………..…………..... 4 2.1. The Importance of ants ………………………………………… 4 2.2. Family Formicidae 5 ……………………………………………. 2.2.1. Main Differential Anatomical Features of 5 Formicidae…….. 2.2.2. …………………………………………….…… 10 2.2.3. The Classification and Origin 10 ……………………………… 2.2.4. Subfamilies 11 …………………………………………………

IV 2.3. The Ants of Sudan 15 ……………………………………………… 2.3.1. History and Geography 15 ………………………………….. 2.4. Genera 18 …………………………………………………………. 2.4.1. …………………………………………….. 18 2.4.2. 18 ……………………………………………… 2.4.3. 18 Monomorium……………………………………………………… 2.4.3.1. Monomorium bicolor Emery 19 (1877)………………………. 2.4.3.2. Monomorium areniphilium Santschi 19 (1911)…………….. 2.4.3.3. Monomorium destrctor Jerdon (1851) ……………….. 19 2.4.3.4. Monomorium mayri Forel (1902) …………………….. 19 2.4.3.5. Monomorium salomonis Linnaeus (1758) ……………. 20 2.4.4. Pheidole …………………………………………………… 20 2.4.5. Tetramorium …………………………………………………….. 20 2.4.5.1. Tetramorium sericeiventris Emery 21 (1877)………….…….. 2.4.6. Camponotus …………………………………………….….. 21 2.4.6.1.Camponotus (Tanaemyrmex) maculates Fabricius (1781) 21 2.4.6.2.Camponotus (Orthonotomyrmex)sericeus Fabricius 21 (1798)………………………………………………… …

V 2.4.7. Cataglyphis 22 ………………………………………………………. 2.4.8. 22 …………………………………………………………. 2.4.9. Paratrechina 22 …………………………………………………….. 2.4.10. Anochetus 22 ………………………………………………………. 2.4.1.1. ………………….………………………… 23 2.5. 23 Karyotyping……………………………………………………… 2.5.1. Chromosomes 23 ……………………………………………… 2.5.2. Somatic chromosomes 24 …………………………………….. 2.5.3. Karyotypes ………………………………………………… 24 2.5.3.1. Karyotpye Study of Ants 24 Chromosomes………………... 2.5.3.2. Karyotpye study of Camponotus 24 …………………. Chapter 3: Material and Methods…………………..………………… 30 3.1. Collection 30 Sites……………………………………………...……

VI 3.2. Specimen collection 32 ……………………………………….……. 3.3. Key used in identification 32 ………………………………..……… 3.4. Karyotyping 32 …………………………………………….……….. Chapter 4: 34 Results……………………………………….…..…………. 4.1. Subfamily: 37 …………………………………….…… 4.1.1. Genus: Cardiocondyla ………………..…………………… 37 4.1.1.1. Characteristics 37 …………………………………….……. 4.1.1.2. Cardiocondyla emeryi Forel (1881) 37 ……………………. 4.1.1.3. Cardiocondyla elegans Emery (1969) 37 ……………….… 4.1.2. Genus: Crematogaster ……... 39 ……………….…………….. 4.1.2.1. Characteristics 39 ………………………………………………. 4.1.2.2. Crematogaster mimosae Santschi 1914 39 …………….….. 4.1.3. Genus: Monomorium 39 ………………………………………. 4.1.3.1. Characteristics 39 ………………………………………………. 4.1.3.2. Monomorium bicolor Emery (1877) 39

VII …………………… 4.1.3.3. Monomorium areniphilum Santschi 41 (1911)…………..… 4.1.3.4. Monomorium destructor Jerdon 41 (1851)…………….….. 4.1.3.5. Monomorium mayri Forel (1902) 41 ………………….….. 4.1.3.6. Monomorium salomonis Linnaeus (1758) 41 …………….. 4.1.4. Genus: Pheidole 41 ……………………………………….…… 4.1.4.1.Characteristics 41 ……………………………………………….. 4.1.4.2. Pheidole sinaitica Mayr (1862) 42 ……………………….. 4.1.4.3. Pheidole pallidula Nylander (1849) 42 ……………………. 4.1.4.4. Pheidole indica Mayr 42 1879………………….………… 4.1.5.Genus: Tetramorium 44 ……………………………….……….. 4.1.5.1.Characteristics 44 ……………………………………………….. 4.1.5.2. Tetramorium sericeventre Emery 44 (1877)……………….. 4.2. Subfamily: 44 ……………………………………..…… 4.2.1. Genus: 44 Camponotus……………………………………….……..

VIII 4.2.1.1.Characteristics 44 ………………………………………………. 4.2.1.2. Camponotus oasium Forel (1890)……………………… 45 4.2.1.3.Camponotus (Tanaemyrmex) maculatus Fabricius 45 (1781). ………………………………………………… 4.2.1.4.Camponotus (Orthonomyrmex)sericeus Fabricius 45 (1798). 4.2.1.5. Camponotus (Myrmopiromis) niveoetosus Mayr (1862). 45 4.2.1.6. Camponotus species 45 5.………………………………….. 4.2.1.7. Camponotus species 6. 45 …………………………………. 4.2.2. Genus: 47 Cataglyphis…………………………………………. 4.2.2.1. Characteristics ………………………………………….. 47 4.2.2.2. Cataglyphis desertorum Forel 47 (1894)……………………... 4.2.2.3. Cataglphis lividus Andre (1881) …………….………… 47 4.2.3. Genus: Lepisiota………………………………………………… 47 4.2.3.1. 47 Characteristics……………………………………………….. 4.2.3.2. Lepisiota gracilicornis Forel 1892 48 …………………….. 4.2.3.3. Lepisiota semenovi Ruzsky (1905) ……………………. 48 4.2.4. Genus: Paratrechina 48 ………………………..……………… 4.2.4.1. Characteristics ………………………………………….. 48 4.2.4.2. Paratrechina longicornis Latreille (1802) ………..……. 48 4.3. Subfamily: 51

IX ……………………………………………. 4.3.1. Genus: Anochetus 51 ……………………………………………….. 4.3.1.1. Characteristics …………………………………………. 51 4.3.1.2. Anochetus traegaordhi Mayr (1904) 51 …………………… 4.3.2. Genus: 51 Pachycondyla……………………………………………. 4.3.2.2. Pachycondyla sennaarensis Mayr 52 (1862)………………. 4.4. Karyotypes of Camponotus (Myrmopiromis) niveoetosus and 52 Camponotus species 6 ………………………………………… 4.4.1.Camponotus (Myrmopiromis) niveoetosus 52 ………………… 4.4.2. Camponotus sp6 …………………………………………. 52 Chapter 5: Discussion ………………..……………..……………….. 56 References ………………………………..…………………………… 58 Appendices ……………………………………..…………………….. 66

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List of Tables Table Page Table I: Imatong Mountain Ants…………………………….. 17 Table II: Species of Ants Recorded From Northern Sudan…… 26 Table III: GPS reading of collection sites …………………….. 33 Table IV: Species of Ants found in Khartoum 34 State…………… List of Figures Figure Page Figure 1: Main Features of the Head of Formicidae ……………. 6 Figure. 2: Anatomical Features of a Generalized …………… 7 Figure. 3: Anatomical Features of a Generalized Ant …………… 8 Figure. 4: Anatomical Features of a Generalized Ant……………. 9 Figure (5): Genus: Cardiocondyla ……………………………… 37 Figure (6): Genus: Crematogaster ………………………………. 39 Figure (7): Genus: Monomorium ………………………………… 39 Figure (8): Genus: Phediole ……………………………………... 42 Figure (9): Genus: Tertamorium ………………………………… 45 Figure (10): Genus: Camponotus ……………………………….. 45 Figure (11): Genus: Cataglyphis ………………………………… 48 Figure (12): Genus : Lepisiota…………………………………… 48 Figure (13): Genus : Paratrechina ……………………………… 49 Figure (14): Genus: Anochetus………………………………...... 52 Figure (15): Genus : Pachycondyla……………………………. 53 Figure (16 a): A spread metaphase plate. The number of 53 chromosomes was 2n=21……………………… Figure (16 b): An Idiogram of Somatic Chromosomes of 53 Camponotus niveoetosus arranged in pairs……. Figure (17 a): A spread metaphase plate. The number of 54 chromosomes was 2n=25

XI Figure (17b): An Idiogram of Somatic Chromosomes of 54 Camponotus species6 arranged in pairs…….

XII List of Plates Plate (I): Acacia fistula Tree………………………………

Plate (Π): Nest of Aphenomyrmex sp………………………………………

Plate (III): Map Showing Location of Khartoum State………………

Plate (IV): Satellite Map of Khartoum State Showing Collection Sites.

Plate (V): Cadiocondyla emeryi…………………………………………... Plate (VI): Cardiocondyla elegans…………………………………..

Plate (VII): Crematogaster mimosae………………………………..

Plate (VIII): Monomorium bicolor……………………………………….. Plate (IX): Monomorium areniphlium ……………………………….

Plate (X): Monomorium destructor …………………………………

Plate (XI): Monomorium mayri …………………………………..… Plate (XII): Monomorium salomonis ………………………………

Plate (XIII): Phediole sinaitica the big headed ant ( Major worker)... Plate (XIV): Phediole sinitica (Minor worker)………………………

Plate (XV): Phediole pallidula ………………………………………..

Plate (XVI): Phediole indica ………………………………………..

Plate (XVII): Tetramorium sericeventre ………………………………... Plate (XVIII): Camponotus oasium …………………………………

Plate (XIX): Camponotus (Tanaemyrmex) maculates ………………

Plate (XX): Camponotus (Orthonomyrmex) sericeus ……………….

Plate (XXI): Camponotus (Myrmopiromis) niveoetosus ……………

Plate (XXII): Camponotus species 5 ……………………………….

Plate (XXIII): Camponotus species 6 ………………………………

XIII Plate (XXIV): Cataglyphis desertorum (sun dog) ……………………

Plate (XXV): Cataglyphis lividus …………………………………….

Plate (XXVI): Lepisiota gracilicornis …………………………….

Plate (XXVII): Lepisiota semenovi ……………………………….

Plate (XXVIII): Paratrechina longicornis ……………………….

Plate (XXIX): Anochetus traegaordhi ……………………………..

Plate (XXX): Pachycondyla sennaarensis …………………………..

XIV List of Appendices ِ Appendix Page Appendix 1: Letters Indicating Collection sites of ants in 66 Khartoum State……………………………………

Appendix 2: Data about collection sites and species collected in 67 Khartoum State……………………………………..

Appendix 3: Arabic Abstract...... 72

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Abstract

Twenty-five species of ants were collected from various regions of Khartoum State. The species were classified into three subfamilies and eleven genera. Of the twenty-five species collected 23 were identified to the species level. Two genera, which included 3 species, were recorded for the first time in Khartoum.

Identification of genera and species depended on morphological characteristics. The identification was confirmed by specialists in Britain and in the of America.

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CHAPTER ONE

INTRODUCTION

Ants are a well-known group among the large number of various on the earth. This is possibly because they are usually quite conspicuous and closely associated with man in his home, garden or place of work. Ants are found in all terrestrial habitats from sub arctic tundra to equatorial rainforest and from deep in the soil to the tips of the highest trees. Their external morphology is more or less uniform except for specialized for feeding or defense purposes. In body size they range from less than one millimeter to about forty millimeters in length.

There are about 9.000 to 10.000 scientifically described ant species. Fisher, (2005) stated that there are 11700 described ant species. All these species are members of a single family: the Formicidae of the Order (Bolton, 1994). Ants are currently classified into 21 subfamilies and 283 genera (Bolton 2003).

The Afro tropical or the Ethiopian region include all sub-Saharan Africa and the southern half of the Arabian Peninsula and excluding and its nearby islands. There are 89 ant genera in the Afro tropical region (Bolton 1994).

1.1. Social Organization

All ant species are social insects that live in colonies in which the number of individuals in a colony ranges from a few dozen to many millions depending on the species (Bolton 1994). The individuals in an consist of three forms or castes:

• A single winged fertile female usually referred to as the queen and whose main function is to lay eggs.

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• A large number of infertile wingless females known as the workers who carry out all the functions of the colony.

• A few winged males whose main function is to fertilize virgin queens during reproductive flights before the establishment of new colonies.

New ant colonies are formed at a certain time of the year, usually during the rainy season. Winged males and females are produced in the established old colonies and usually fly out of the colony at a certain time. The flight time is usually synchronized with that of similar individuals of the same species from other colonies found in the area. Each individual pairs with an individual of the opposite sex usually from another colony. Synchronization of the flight of the winged males and females prevents inbreeding within the same colony. The pair mate, the female lays eggs and the male usually dies. The eggs hatch into larvae, which develop into wingless workers that start the building of a new colony. Nests can be simple cavities in the soil, a space beneath a piece of bark or in some species leaf shelters stitched together with larval silk. Certain species build large external earth mounds with complex subterranean excavations. Some species form symbiotic associations with certain plants such as Pseudomyrmex species with Acacia fistula trees Plate 1 and Plate II.

1.2. Feeding

Various species feed on plant seeds, nectar, honey-dew secreted by sap-sucking insects, and fungi, but most are general or specialized carnivores that can feed mainly on other invertebrate . A few are specialized in preying on other ants. Some are scavengers that feed on dead and plant material. Various ant species have specialized behavioural adaptations such as culture of fungi, seed harvesting, herding

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Plate (I): Acacia fistula Tree

Plate (Π): Nest of Pesudomyrmex sp

and milking of aphids and other plant sucking bugs and social parasitism or slave making.

1.3. Objectives of this study

Sudan is a huge country with highly diverse habitats ranging from desert in the north, semi desert, poor savanna, rich savanna, and tropical rain forest in the extreme south. This diversity in geographical habitats led to a large diversity in species of animals and plants. Ant fauna has not been studied in the Sudan. However, Weber (1943c) recorded and described 125 species from the Imatong Mountains in the Southern Sudan. In this work a survey of ant species in Khartoum State has been carried out with a view of identifying the species and learning some of the techniques used in ant taxonomy.

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CHAPTER TWO

LITERATURE REVIEW

2.1. The Importance of ants

Ant species constitute only 1.5 % of the known global fauna but they make up about 10% of the total animal biomass in tropical forests, grasslands and probably other major habitats. They are among the major predators of as well as important scavengers in most terrestrial habitats. They turn and enrich more soil than earthworms Wilson, (1971), Jeanne, (1979), Levieux, (1982) Sorensen and Schmidt, (1987)). Thus ants have an important impact on the terrestrial environment.

Leafcutter ants such as members of the genera Acromyrmex and Atta, are the principal herbivores and the most destructive insect pests of Central and South America (Weber, 1972, Cherrett, 1982). Pogonomyrmex and other harvester ants are principal granivores, competing effectively with mammals for seeds in deserts of the Southwestern United States (Davidson et al, 1980).

In certain regions ants are responsible for the dispersal of seeds of many herbaceous plant species (Handel et al, 1981). They aid in the spread of forest vegetation onto bare rocks in Finland (Oinonen, 1956) and dune vegetation on to salt lakes in the USSR (Pavlova, 1977).

The abundance and ecological dominance of ants are matched by their extremely large geographical range. The only places free of native ant species are Antarctica, Iceland, Greenland, Polynesia east of Tango, and a few of the remote islands in the Atlantic and Indian Oceans (Wilson and Taylor, 1967b). Four genera (Camponotus, Crematogaster, Hypoponera, and Phediole) are found all over the world ( Wilson,1976e).

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2.2. Family Formicidae:

Formicidae are social Hymenoptera with a wingless female worker caste and winged male and female reproductive castes that live in colonies.

2.2.1. Main Differential Anatomical Features of Formicidae

The head is prognathous in female castes (workers and queens). Antennae have 4 to 12 segments in the female caste and 9 to 13 segments in males. The antennae are elbow shaped bent between the long basal segment and the remaining distal segments (Fig 1).

The second abdominal segment is reduced, forming a node or a scale known as the petiole isolated from the altitrunk in front and the remaining abdominal segments behind (Fig 2), (Fig 3) and (Fig 4).

Frequently the third abdominal segment is also reduced and isolated and known as the post-petiole (Fig 2). Wings of queens are deciduous and usually shed after mating (Bolton 1994).

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Figure 1: Main Features of the Head of Formicidae

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Figure. 2: Morpholgical Features of a Generalized Ant

7

Figure. 3: Morphological Features of a Generalized Ant

8

Figure. 4: Morphologicalii Features of a Generalized Ant

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2.2.2. Taxonomy

Forel (1878) divided the Family Formicidae into five subfamilies: Camponotinae, Dolichoderinae, , Ponerinae, and Myrmicinae. These remained unchanged for many years. However, the Camponotinae had its name changed later to Formicinae by Wheeler (1920). Bolton (1994) lists 16 subfamilies in the Formicidae. The new eleven subfamilies were established by splitting the original 5 subfamilies or by the discovery of new species that constitute subfamilies unknown to the early classifiers.

2.2.3.The Classification and Origin

Known living ants are classified into 11 subfamilies, 303 genera, and possibly 9000 to 11700 species. The number of known species is far below the number that actually exists. This is because the number of nondescribed species is immense especially in the tropics. Overall, it is quite possible that 20,000 or more species of ants, including as many as 350 genera, exist in the world ( Holldobler and Wilson 1990).

The taxonomy of the world ant fauna is still very incomplete. There are few useful regional monographs such as Creighton's (1950) review of the ants of North America and North of . Ant larvae have been described by Wheeler and Wheeler (1951,1986) with a supplementary supplied by Picquet (1958).

The cytotaxonomy of ants, which, is currently a very important subject due to advances on genetics and molecular biology, has been reviewed by Crozier (1975), Imai et al. (1977), and Hauschteck-Jungen (1976,1983).

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2.2.4. Subfamilies

1- Subfamily: AENICTINAE

The single genus (Aenictus) included in this subfamily occurs throughout the Afrotropical, Oriental and Indo-Australian regions. A few species also occur in the southern Palaearctic and northeastern part of the Australasia region. It is absent from the Malagasy, Nearctic, and Neotropical regions.

2- Subfamily: AENICTOGITONINAE

This small subfamily contains only seven species in a single genus (Aenictogiton) known only from its morphologically different males. It is restricted only to Central Africa (Bolton 1994).

3- Subfamily: ANEURETINAE

The single extant genus ( Aneuretellus, Aneuretus, Mianeuretus, Paraneuretus, Protaneuretus) and species of this subfamily is restricted to the island of Sirilanka. Some extinct genera come from the Baltic Amber, Russia, and the U.S.A. (Bolton 1994).

4- Subfamily: APOMYRMINAE

The single genus Apomyrma of this subfamily is represented by two West African species. They have been recorded from , , , , and .

5- Subfamily: CERAPACHYINAE

Two of the five Cerapachyinae genera, Acantostichus and Cylindromyrmex are mostly restricted to the Neotropical region, though the former extends its range into the extreme southern Nearctic; the other three are more widely distributed. Simopone is found in the Afrotropical region, Malagasy, and Indo-Australian regions, with the greatest number

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of species in the Afrotropical region. Sphictomyrmex is predominantly Australian but a few species of this genus, also, occur in the Afro tropical region, Oriental, Indo-Australian, and Neotropical regions. Cerapachys is the largest and most widely distributed genus of the subfamily, with representatives in all Zoogeographical regions. The greatest number of species occurs in the Palaearctic, Nearctic, and Neotropical regions.

6- Subfamily: DOLICHODERINAE

The subfamily Dolichoderinae has a worldwide distribution; its genera are either endemic or artificially introduced by human activities (Bolton 1994).

7- Subfamily: DORLINAE

The single genus ( Dorylus) of this subfamily is found in the southern Palaearctic, Afrotropical, Oriental, and Indo-Australian regions, with the vast majority of species in the Afrotropical region.

8- Subfamily: ECITONINAE

The five genera of this subfamily are distributed in the new world, with the vast majority of species in the Neotropical regions. The genera Cheliomyrmex and Eciton are restricted to this region but a few species of Neivamyrmex. Nomamyrmex, and Labidus occur in the Nearctic region (Bolton 1994).

9- Subfamily: FORMICINAE

This is the second largest subfamily of ants and includes 43 genera and 1400 species, 600 of which are in the genus Camponotus. Formicinae is a cosmopolitan subfamily, which is dominant in temperate regions and common in the tropics. The members of this subfamily are morphologically the most highly developed of all ants. This may be due to the fact that their habits are very diverse. They feed largely on plants

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and plant products. However, many Camponotus species favour sugary substances such as nectar and honeydew, which are sometimes stored in special, replete forms of worker individuals such as in the case of the honey-pot ants Melophorus, and Myrmecocystus (Wheeler and Wheeler 1972).

The nesting habits in this subfamily are very varied. Oecophylla and a few Camponotus species build nests in leaves using their larvae as silk-producing shuttles to sew up the leaves (Wheeler and Wheeler 1972).

Certain of Formica and Polyergus are social parasites or “slave makers”. Polyergu species are true social parasites of Formica. They attack nests of Formica, kill the adults and carry off their pupae to their own nests. The pupae develop into workers which serve and take care of Polyergu adults and offspring (Wheeler 1922a).

10- Subfamily: LEPTANILLINAE

This subfamily otherwise the genus Leptanilla occurs in all the old world regions is entirely absent from the new world and the Malagassy region. There are four genera known only from males and are found in the Indo-Australlian region (Noonilla, Phyaulomyrma, Scyphodon) and the Palaearctic and Oreiental regions (Yavnella). The two remaining genera, Anomalomyrma and Protanilla, are centred on the Indo- Australian region, where the greatest number of species is in the Palaearctic and in the Oriental region.(Bolton 1994).

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11- Subfamily: LEPTANILLOIDINAE

The single genus Leptanilloides of this subfamily is restricted to the Neotropical region. Only two species are known, each from a single collection in Bolivia and .

12- Subfamily:

The single extant genus of this subfamily( , Prionomyrmex, Ameghinoia, Cariridris) is restricted to the Australian region. All the species are from Australia excepet for one, which occurs in New Caledonia.

13- Subfamily: MYRMICINAE

The Myrmicinae are the largest subfamily of Formicidae comprising 2000 species in 155 genera. They are a cosmopolitan group, which is heterogeneous in both anatomy and habits, ranging from primitive to highly specialized species. Among those specialized in feeding habits are the harvester ants (Messor ) and fungus-grower ants (Trachymyrmex). The Myrmicinae also include most of the social parasites which in certain cases have lost their worker caste (Wheeler and Wheeler 1972).

14- Subfamily: NOTHOMYRMECIINAE

The single extant genus () of this small subfamily is found only in the Australasian region, where it occurs in South and West Australia.

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15- Subfamily: PONERINAE

This subfamily is a primitive group of 350 species in 57 genera. The greatest number is found in the southern hemisphere, but 13 genera are found in the Holarctic region. The Ponerinae are entomophagous with usually small colonies. They build small nests in the soil or old logs. The Ponerinae are economically important in the tropical countries as an estimated 80% of their food is (Wheeler and Wheeler 1972).

Distribution

The subfamily ponerinae is found in all zoogeographical regions.

16- Subfamily:

Of the three genera, which make up this subfamily, Myrcidris and Pseudomyrmex are found only in the New World while Tetraponera is very widely distributed in the old world. The genus Myrcidris is known only from , but Pseudomyrmex occurs in all other regions of the New World. Tetraponera is present in all zoogeographical regions of the old world.

2.3. The Ants of Sudan

2.3.1. History and Geographyy

Magretti (1884) collected in eastern Sudan and ; the ants were identified by Ernesto, Andre and Magretti (1884). Magretti described how minute his collection efforts were in an area of about two thousand kilometers. In 1883, the year before the uprising against General Gordon, Magretti traveled from the port of Suakin, collecting in the road to Kassala (on Sudanese side of broder with Eritrea. in the desert plain of Bani Homrani, on the shore at Gasch, on the banks of the Settit at Takazzi (Gasch-Settit Reserve now in Eritrea), and from Bahr el Salaam, ending the remote Metemma in Galabat ( Galabat, -Sudan border).

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Secondly he returned to Kassala, going via Sennehit, with short stops at Karen, Bogos and returning to Massawaa. .He had left Suakin at the end of January and was back there on the first of May.

Even earlier, specimens from an unnamed collector had been deposited in the Museum in Vienna.

Weber (1942a, 1943c) studied ants of the Imatong (or Lolibai) Mountains of South Sudan Table (I). Much of the ant fauna he described belong taxonomically to the Congo Basin faunal group (Taylor 2005)

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Table I: Imatong Mountain Ants

Subfamily No of species

Dorylinae 3

Cerapachyinae 2

Ponerinae 25

Myrmicinae 54

Dolichoderinae 8

Formicinae 33

Total Number 125

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2.4. Genera:

2.4.1. Cardiocondyla

The genus Cardiocondyla contains about 40 species, mostly distributed in the old world (Bolton 1982).

Species of this genus recorded from North Sudan are listed in Table (II)

2.4.2. Crematogaster:

Crematogaster is a very large genus with considerable taxonomic confusion. Bernard (1952) wrote of the tribe Crematogasterini, with its single genus Crematogaster as the most difficult to classify of all the tropical ants. Santschi catalogued the Crematogaster of Congo( Taylor, 2005).

The attachment of the postpetiole to the gaster in this genus allows the characteristic defence posture in which the gaster is turned forwards over the alitrunk.

Many members may have economic importance, especially as several species are among the ants which effectively constitute much of the insect fauna of cocoa and other trees. Members of the genus are omnivores and are the only ants which effectively resisted raids by nomadic Dorylus army ants Bernard (1952).

Species of these genus recorded from North Sudan are listed in Table (II).

2.4.3. Monomorium

Monomorium is a large genus which contains some 300 species half of which occur in the Afrotropical Zoogeographical region. The main distribution reigons of Monomorium include Africa, Australia, with

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secondary centers in the oriental region (Bingham 1903) and Madagascar (Bolton 1987)

2.4.3.1. Monomorium bicolor Emery (1877)

This is the most successful and widely distributed species of its group in Afrotropical region. Monomorium bicolor also extendes it range into the drier zones of Southern Palaearctic, being found in North Africa and . In Sub-Saharn Africa Monomorium bicolor is a species of open Savannah and semi-arid zones, but it also found in forested areas in Nigeria and Ghana. Nests are constructed directly into the earth and the species appears to be a general scavenger in habit, easily collected in traps baited with crushed insects.(Bolton 1987).

2.4.3.2. Monomorium areniphilium Santschi (1911)

Appears to be a successful species found north and south of the . It shows variation in colour and size in its wide ranging habitat (Collingwood and Agosti 1996).

2.4.3.3. Monomorium destructor Jerdon (1851)

Monomorium destructor is most probably a species of Indian origin. It is widely distributed throughout the tropical zones of the World and is increasingly being spread into the temperate zones by commercial activities as it is able to survive in heated buildings. On describing the species Jerdon (1851) noted that these ants prefer animal to vegetable food, destroying dead insects, bird skins, but they also feed on sugar. They are common in all parts of (Bolton 1987).

2.4.3.4. Monomorium mayri Forel (1902)

The range of Monomorium mayri covers a very wide continuous broad band of territory, stretching from east to west of the Indian sub continent. Its range extends westward from there across the middle and

19

near East and through the Sahelian zone of Sub-Saharan Africa. East of India M. mayri was recorded from and Malaysia (Bolton 1987).

2.4.3.5. Monomorium salomonis Linnaeus (1758)

This is one of the largest species of the genus Monomorium and they are distributed throughout Africa south of the Sahara and inhabit dry ground and well-drained soils in Savannah and Semi-desert areas. They nest in exposed places where the soil receives direct sunlight (Bolton 1987).

Species of this genus recorded from North Sudan are listed in Table (II).

2.4.4. Pheidole

This is a very large genus, with variable nesting sites and foraging habits. Bernard (1952) noted there were 76 species known from Africa; of which 44 were from French West Africa. The classification was complicated by the presence of workers and soldiers, with enormous heads. Because of the relative similarity of males, queens and workers, it was found that the soldiers were useful for distinguishing species. Taylor (2005) found the soldiers, in general, to be much more useful in distinguishing the various species.

Species of this genus recorded from North Sudan are listed in Table (II).

2.4.5. Tetramorium

In this genus erect hairs were simple in most species, but were divided in many former Triglyphothrix, and bizarre in a few species. The former Macromischoides (or at least African members) do not have the posterior lateral margins of the clypeus raised into a ridge bordering the antennal insertions; are less noticeably square- shouldered; and have a

20

more rounded shape to the head in full face view, plus antennal scapes which surpass the occipital margin(Taylor 2005).

Species of this genus recorded from North Sudan are listed in Table (II).

2.4.5.1. Tetramorium sericeiventris Emery (1877)

This ant is an ant of dry plains north of the Imatong Mountains. In Khartoum it had the curious habit of using the feces of small birds, especially the Sudanese race of Passer domesticus, on its burrows probably to act as a barrier against the sand of the desert which tended to fill the nest entrance (Weber 1943).

2.4.6. Camponotus

This is the biggest ant-genus distributed all over the world. Camponotus includes 500 species and many subspecies (Bernard 1952). Taylor(2005) reported 145 species from Africa.

2.4.6.1.Camponotus(Tanaemyrmex) maculates Fabricius(1781)

This common African species was also found in the southern warmer valleys of Arabia (Collingwood, and Agosti 1996)

2.4.6.2.Camponotus (Orthonotomyrmex)sericeus Fabricius(1798)

This ant is large and conspicuous and forages throughout the day. (Collingwood and Agosti 1996).

Species of this genus recorded from North Sudan are listed in Table (II).

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2.4.7. Cataglyphis

The genus Cataglyphis is characterized by well-diagnosed monophyletic groups based on male genitalia characters but the species- level taxonomy is hardly understood ( Agosti 1990).

Cataglyphis species are distributed in the Palaearctic from Portugal to Mongolia and the Ganges valley in India and from the Ivory Coast and Southern Sudan northwards to Czechoslovakia (Agosti 1990).

Species of this genus recorded from North Sudan are listed in Table (II).

2.4.8. Lepisiota

Strickland (1951a) described members of this genus as small black, soil nesting ants (Taylor 2005).

Species of this genus recorded from North Sudan are listed in Table (II).

2.4.9. Paratrechina

Bernard (1952) described members of this as small, very specialized, terricolous ants, showing polymorphism. The larvae have reduced mouthparts and are fed on the exudates of Homopetra, which these ant workers often raised in their nests (Taylor2005).

Species of this genus recorded from Northern Sudan are listed in Table (II).

2.4.10. Anochetus

The colony size of this genus is small usually about 100 individulas. They usually nest in rotten, hallow twigs, or under bark. They are probably all predators that mostly forage after dark and use a

22

trap-jaw mechanism to seize prey (Brown 1976). They are usually found in the tropics (Taylor 2005).

Species of this genus recorded from Northern Sudan are shown Table (II).

2.4.11. Pachycondyla

This genus is a large group of ants with about 200 described species mostly known from the tropics and sub-tropics ( Bolton, 1995). The species of Pachycondyla are diverse in their morphology (Wild, 2002). Most species appear to be general scavengers and predators of arthropods, which they subdue with venom (Orivel and Dejean 2001).

The nest and social biology of Pachycondyla is variable. They live in colonies of a few dozen to a few thousand workers and nest in a variety of conditions, including soil, leaf litter, rotting wood, dead branches, plant cavities, and epiphytes (Leal and Oliveira 1995, Longino, 2002, Lucas et al 2002, Trunzer et al 1998, Wheeler 1972, Wild, 2002).

Species of this genus recorded from North Sudan are listed in Table (II).

2.5. Karyotyping

2.5.1. Chromosomes

Chromosomes are very important tools in the classification and taxonomy of animals and plants (Fristrom and Clegg, 1988; El Banhawy et.al.1994). Different species possess different numbers of chromosomes and individuals of the same species contain the same number of chromosomes in their cells (Schjede and De vellis, 1970; Fristorm; and Clegg; 1988; El Banhawy et .al; 1994). Chromosomes are divided into two groups: somatic and sex chromosomes.

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2.5.2. Somatic chromosomes

Somatic chromosomes usually occur in a double number or diploid set (2n). Chromosomes are morphologically identified by two features; the relative size and the position of the centomere (Swanson, 1957, Fristorm and Clegg, 1988, Suzuki; et. al, 1986; El Banhawy et. al., 1994).

2.5.3. Karyotypes

A karyotype means the group of characteristics that may be taken into consideration when identifying the chromosomal complex of a species. These characteristics include shape, length and diameter of the chromosomes, and position of their centromeres. The karyotype is characteristic for each species and is usually represented by a diagram known as the ideogram (Swanson 1957; El Banhawy et.al., 1994). The karyotype can either be designated in haploid (n) or diploid (2n) number (Badawi 2002).

2.5.3.1. Karyotpye Study of Ants Chromosomes

Karotypes of ants of the subfamily Formicinae of India were studied by Imai et al (1984) and were found to be characterized by highly divergent chromosome numbers: 2n =16, 18, 20, 24, 28, 30, 34, 35, 39, 40,42,44 and 45.

2.5.3.2. Karyotpye study of Genus Camponotus

This genus is one of the most successful ant genera distributed all over the world and includes more than 500 species (Imai et. al.1984).

This ant genus is as karyotypically heterogeneous as its species are morphologically different. The known chromosome numbers range between 2n= 18 to 50 (Crozier 1975). Rather divergent chromosome numbers were also found among Indian Camponotus. The numbers

24

recorded for species of this genus were 2n=20, 24,26,32,34,35,39,40 and 44 ( Imai et al.1984).

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Table II: Species of Ants Recorded From Northern Sudan.

Year of Species Author Location of finding publication Cardiocondyla emeryi mahadii Karavaiev 1911 Khartoum Cardiocondyla emeryi mahadii Weber 1952 Port Sudan Cardiocondyal nilotica Weber 1952 Ed Dueim (Abduwaym), White Nile Crematogaster(cr.) rectangulata Viehmeyer 1923 Port Sudan Crematogaster(cr.) aegyptiaca Viehmeyer 1923 Port Sudan Crematogaster(cr.) aegyptiaca Viehmeyer 1923 Sennar, Elobied in hills, robusta Bara, Kaduugli Crematogaster(cr.) anataris or Viehmeyer 1923 Elobeid, Bara jehovae Crematogaster(cr.) chiarinii Santschi 1914a subsulcata Crematogaster(cr.) chiarinii Viehmeyer 1923 Sennar and Talodi subsulcata Crematogaster(cr.) senegalensis Andre 1884 Suakin, Metamma, Sebderat Crematogaster(cr.) senegalensis Mayr 1904 Omdurman Crematogaster(cr.) senegalensis Viehmeyer 1923 Nubba ,kaka-Sungikai Crematogaster(cr.) Mayr 1895 sewellimarnoi Crematogaster(cr.) Viehmeyer 1923 Tongo sewellimarnoi Crematogaster(spaer) Viehmeyer 1923 Dilling gambiensis Crematogaster(spaer) Santschi 1914 Lado, Redjaf gambiensis transersiruga Crematogaster(spaer) Viehmeyer 1923 Nubba, kaka, Gulfan, Keiga, pseudinermis Tummero Monomorium abyssinicum Viehmeyer 1923 Dilling M. afrum Andre 1884 Atbara M. afrum asmarensis Viehmeyer 1923 Tongo M. barabatulum Andre 1884 M. bicolor Andre 1884 Suakin, Metamma M. clavicorne Mayr 1904 Kaka M. destructor gracillium Andre 1884 Kassala

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M. hercules Viehmeyer 1923 Elobeid bis Bara,Atbara M. mayri karawajewi karavaiev 1911 Khartoum M. mayri karawajewi Viehmeyer 1923 Khartoum M. nitidiventre Viehmeyer 1923 Nubba, kaka-Sungikai Keigatummero, Kadugli M. osscaris despecta Viehmeyer 1923 Tanguru M. pharaonis Andre 1884 M. pharaonis Viehmeyer 1923 On Nile steamer "Omdurman"

M. salomonis Andre 1884 M. salomonis Finiz 1939a Port Sudan M. salomonis Mayr 1904 Omdurman Phediole aeberlii Mayr 1904 Omdurman Phediole megacephala Mayr 1904 Abba island Phediole megacephala Viehmeyer 1923 Elobeid impressifrons Phediole rugaticeps Andre 1884 El Hefera, Setti Phediole rugaticeps arabs Viehmeyer 1923 Elobeid, UmmRamad, Nubba, kaka, Korokak Phediole sinaitica Andre 1884 Atbra River, Sebderat Phediole speculifera Andre 1884 Bahr-el-Salam Phediole termitophila Forel 1904 Kaka

Camponotus(Myrmopiromis) Andre 1884 niveosetosus Camponotus(Myrmosericus) Viehmeyer 1923 Sennar cinctellus Camponotus(Myromsericus) Viehmeyer 1923 Dilling and Raggaba flavomarginatus Camponotus(Myrmotrema) galla Santschi 1933 Khatoum Camponotus(Myrmotrema) galla Viehmeyer 1923 Khartom and Sennar Camponotus(ortho) sericeus Santschi 1933 Khartoum and Shendi obtusus Camponotus (Tanae) fellah Viehmeyer 1923 Port Sudan Camponotus (Orthoon.) sericeus Andre 1884 Kor lebka Camponotus(orthon.) sericeus Viehmeyer 1923 Bara,Dilling and Tanguru.

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Camponotus(Tanae.) maculatus Andre 1884 Camponotus(Tanae.) maculatus Viehmeyer 1923 Many places. Camponotus(Tanae.) maculates Andre 1884 Suakin, Kor langhebb congnatus Camponotus(Myrmotrema) Viehmeyer 1923 Sennar diplopunctatus subconvexus Cataglyphis bicolor Viehmeyer 1923 Many places Cataglyphis bicolor sudanicus Karavaiev 1911 Cataglyphis bicolor sundanicus Santschi 1933 Kodraco Cataglyphis bicolor/ viaticus Andre 1884 Sebderat and Kassala. Cataglyphis bombycinus Viehmeyer 1923 Wadi Halfa Cataglyphis viaticus desertoum Mayr 1904 Gebelein Cataglyphis lividus Viehmeyer 1923 Elobeid-Bara Lepisiota canescens Mayr 1904 Kaka Lepisiota canescens Viehmeyer 1923 Sennar, Elobied, Karshawal Andre 1884 Sebdrat near Kassala Lepisiota frauenfeldi Andre 1884 Lepisiota gracilicornis Viehmeyer 1923 Gulfan, Aswan, Portsudan Lepisiota obtusa Firizi 1939a Portsudan. Paratrechina longicornis Viehmeyer 1923 Portsudan Paratrechina traegaorcdhi Forel 1904 Kaka. Anochetus traegeordhi Mayr 1904 Khartoum Anochetus traegeordhi Viehmeyer 1923 Renk gracilicornis Anochetus sedilloti Brown 1978 Khor, near Umm Dorein Pachycondyla(Brachy.) Andre 1884 Bahr-el-Salaam Kassla sennaarensis Pachycondyla(Brachy.) Mayr 1862 Sennar sennaarensis Pachycondyla(Brachy.) Mayr 1904 Khartoum sennaarensis Pachycondyla(Brachy.) Santschi 1933 Khartoum sennaarensis Pachycondyla(Brachy.) Viehmeyer 1923 Several locations sennaarensis Pachycondyla(Brachy.) Santschi 1937 Khartoum sennaarensis

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Pachycondyla(Megaponera) Andre 1884 Sennar analis Pachycondyla(Megaponera) Viehmeyer 1923 Several locations analis Pachycondyla(Paltothyreus) Viehmeyer 1923 Dilling tarsatus Tetramorium sericeiventre Andre 1884 El Hefera, Settit Tetramorium sericeiventre hori Santschi 1818b Khartoum Tetramorium simillimum Mayr 1904 Omdurman Aenictus hamifer Santschi 1933 Khartoum Cataulacus intrudents hararicus Viehmeyer 1923 On steamer Omdurman Dorylus fimbriatus laevipodex Santschi 1933 Khartoum Leptothorax angulatus Bolton 1982 Khartoum-PortSudan Messorbarbarus Andre 1884 Several locations Messor galla Karavaiev 1912 Khartoum

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Plate (III): Map Showing Location of Khartoum State.

CHAPTER THREE

MATERIALS AND METHODS

3.1. Collection Sites

Ant specimens were collected during the period April to October 2004 from different localities in Khartoum State. The study sites are shown in Plate (Ш) and Plate (IV).

Khartoum State has an average daily maximum and minimum temperatures of 39 - 26° c in summer and 34°c - 19°c in winter. An average relative humidity of 29 % and an annual rain fall of 30 mm from July to November (Data from Khartoum Meteorological Station).

3.2. Specimen collection

Ants were mostly collected manually usually from nests or foraging columns. Sometimes a moistened paintbrush, a pair of forceps or an aspirator was used for collection. The specimens were stored in plastic or glass tubes containing 70%-75% ethanol. The time of collection was during the early morning and in the mid of the day.

3.3. Key used in identification

Bolton’s key (1994) was used for identification of the specimens up to the generic level.

3.4. Karyotyping

Chromosome preparations were made from the brains of workers of Camponotus niveoetosus and Camponotus species 6.

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N #Shambat

Umbada # Tuti# Island Cokoo# River# Bank #Khartoum University Botonical Gaden # Ac# aci a f or es t

Sal# ha

Rawk keb #

Soba# Farm

Plate (IV): Satellite Map of Khartoum State Showing Collection Sites.

Chromosome spreads were prepared by applying a modification of the karyotyping technique of Baker (1970) wich has been modified by (Badowi 2002). Tissues were dissected out and were put in 3 ml of freshly prepared 0.5% potassium chloride (KCL) hypotonic solution (0.5 gm KCL +100 ml distilled water). A glass rod was used to break the cell clumps. The resultant suspension was allowed to settle for 15-20 minutes. The cell suspension was then centrifuged at 300 rounds per minute for ten minutes, and the supernatant fluid was discarded leaving the settled undisturbed cells at the bottom of the tube. Three milliliters of freshly prepared Carony’s fixative were added and the settled cells were mixed gently and were allowed to fix for ten to fifteen minutes. The cells were centrifuged again for five minutes at 300 rounds per minute. The supernatant fluid was discarded leaving a small quantity over the settled cells. The cells were resuspended in one milliliter of fresh fixative and centrifuged for five minutes. The last step was repeated three times. Finally, the cells were stored in the fixative in a test tube.

The slide was placed on a warm plate and two to three drops of the fixed cell suspension were placed on the slide using a dropper and the slide was then left to dry. The slide was stained in Giemsa’s stain (one part Giemsa’s stock solution + three parts distilled water) for ten to fifteen minutes, and then washed with distilled water.

The slides were left in open air for one day to dry and were then examined under the microscope and suitable specimens were photographed with the help of a phase contrast microscope using an oil immersion lens.

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CHAPTER FOUR

RESULTS

Twenty-five species of ants were collected from Khartoum State. They were classified into three subfamilies:

1- The Myrmicinae with 12 species.

2-The Formicinae with 11 species

3- The Ponerinae with 2 species.

The species were grouped in eleven genera and were identified using individuals from the worker caste, according to the key of Bolton (1994).The major characteristics of each genus and species were recorded. Fig. (5) to Fig. (15) and Plate (V) to Plate (XXX) shows the main anatomical characteristics on which the description depended.The distribution of the various species in Khartoum State is shown in Table (IV). The GPS ( Global Position System) readings of collections sites are shown in table (III).

Table shown in Appendix (2) shows data and information about localities of ants collected during this work. Information in this table were sent to Dr. Brian Fisher, California Academy of Sciences on his request to be published them in the Academy website www.antweb.org as Ants of Sudan.

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Table III GPS Readings of Collection Sites

Locality Latitude Longitude Elevation Botanical Gradern 15° 36.066' 32° 30.572' 396m Acacia Forest 15° 35.895' 32° 29.880' 380m Shambat 15° 40.360' 32° 31.520' 392m River Bank 15° 36.789' 32° 32.366' 379m Salha 15° 33.375' 32° 27.127' 410m Umbada 15° 38.885' 32° 27.599' 416m University of Khartoum 15° 36.643' 32° 32.500' 402m kukuo 15° 37.111' 32° 34.909' 396m Soba Farm 15° 22.631' 32° 39.725' 400m 15° 2' 32º 0' 420m Rawkeeb 15° 36' 32º 10' Tuti island 15° 36' 32° 30' 425m

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Table IV: Species of Ants found in Khartoum State. Locality Subfamily and Species River Acacia Botonical Soba Tuti Khartoum Kokoo Shambat Rawakeeb Salha Umbda Bank Forest Garden Farm Iland University ٭ ٭ Subfamily: Ponerinae ٭ ٭ ٭ Anochetus traegaordhi ٭ ٭ ٭ ٭ ٭ ٭ ٭ ٭ ٭ Pachycondyla sennaarensis ٭ ٭ ٭ ٭ Subfamily: Formicinae ٭ ٭ ٭ ٭ ٭ ٭ ٭ ٭ ٭ Camponotus oasium

٭ ٭ Camponotus maculatus ٭ ٭ ٭ ٭

٭ ٭ Camponotus sericeus ٭ ٭ ٭ ٭ ٭ Camponotus niveosetosus ٭ ٭ ٭ ٭ Camponotus sp5 ٭ ٭ ٭ Camponotus sp6 ٭ ٭ ٭ Cataglyphis desertorum ٭ ٭ ٭ ٭ Cataglyphis lividus ٭ Lepisiota semenovi ٭ Lepisiota gracilicornis ٭ Paratrechina longricornis

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٭

٭

٭

٭ ٭ ٭

٭ ٭ ٭ ٭

٭

٭ Subfamily: Myrmicinae ٭ ٭

Cardiocondyla emeryi

Cardiocondyla elegans

٭ Crematogaster mimasoe ٭ ٭

٭ ٭ ٭ Monomorium bicolor ٭ ٭ ٭ ٭ ٭ Monomorium areniphilum ٭ ٭ Monomorium destructor ٭ Monomorium salomonis ٭ Monomorium mayri

Tetramorium sericeventre

Pheidole pallidula

٭ Pheidole sinatica(major worker) ٭

Pheidole sinatica (minor worker)

Pheidole indica

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Plate (V): Cardiocondyla emeryi

Plate (VI): Cardiocondyla elegans

4.1. Subfamily: Myrmicinae

4.1.1. Genus: Cardiocondyla Fig (5).

4.1.1.1. Characteristics

1- The antennae were formed of 12 segments each Fig (5).

2- The propodeal lobes were low and rounded. The propodeum was usually armed with cuticular teeth or spines and only extremely rarely unarmed. When there are propdeal spines, they were not connected to the lobes Fig (5).

Two species of this genus were found in Khartoum Sunt Forest on the right bank of the White Nile. They were extremely small ants usually less than one millimeter in length. Both species were found in rotten logs of Acacia nilotica.

4.1.1.2. Cardiocondyla emeryi Forel(1881)

In this species the petiole in profile had a flattened dorsum. The colour was orange and the gaster and the antennal club were brown. Plate (V).

4.1.1.3. Cardiocondyla elegans Emery 1969

The alitrunk, peduncle, and gaster were shiny. Propodeam was strongly built and postpetiole was wider than the petiole. The overall colour was dull brown Plate (VI).

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Figure (5): Genus: Cardiocondyla

37

Plate (VII): Crematogaster mimosae

Plate (VIII): Monomorium bicolor

4.1.2. Genus Crematogaster Fig (6)

4.1.2.1. Characteristics

1-The postpetiole was articulated on the dorsal surface of the first gaster segment.

2-The gaster when seen from the dorsal side was roughly heart- shaped and capable of reflexion over the alitrunk.

3-The petiole was dorsoventrally flattened and without a node.

4.1.2.2. Crematogaster mimosae Santschi 1914

Head and alitrunk strongly sculptured spines broadly dentate. Whole body coloured reddish brown Plate (VII).

The species was found in a tunnel in a stem of Acacia nilotica tree.

4.1.3. Genus: Monomorium Fig (7)

4.1.3.1. Characteristics

1- The median portion of clypeus was distinctly raised.

2- The postpetiole node was less voluminous than the petiole node in profile and was narrowly attached to the gaster.

3- Propodeum was usually unarmed.

4.1.3.2. Monomorium bicolor Emery (1877)

The dorsal part of the propodeam had a slight shallow cavity. The head and alitrunk were red to bright orange-yellow in colour and the gaster was black Plate (VIII). This species was very common in houses, stores and farms and considered as a domestic pest.

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Figure (6): Genus: Crematogaster

Figure (7): Genus: Monomorium

39

Plate (IX): Monomorium areniphlium

Plate (X): Monomorium destructor

Plate (XI): Monomorium mayri

Plate (XII): Monomorium salomonis

4.1.3.3. Monomorium areniphilum Santschi (1911)

The ventral aspect of the head had always a few projecting hairs Plate (IX).

This species had the second largest distribution in Khartoum State and is considered as pest in houses, stores and farms.

4.1.3.4. Monomorium destructor Jerdon (1851)

Anterior body from head to postpetiole had a yellowish brown or reddish brown colour. The gaster was blackish brown Plate (X).

Petiole node in dorsal view subglobular ; postpetiole node only slightly broader than long.

4.1.3.5. Monomorium mayri Forel (1902)

The occiput as well as the propodeum have fine transverse striations. Anterior body coloured brown, gaster black Plate (XI).

4.1.3.6. Monomorium salomonis Linnaeus (1758)

The mesopropodeal furrow was distinct, making a right angle in profile Plate (XII). Anterior body was shiny yellow and gaster was shiny black.

4.1.4. Genus: Pheidole Fig (8)

4.1.4.1.Characteristics

1- In this genus the terminated in a strongly defined three segmented club.

2- The pronotum or pronotum and anterior mesonotum formed a high dome or a markedly convex arc. The propodeum had a separate convex or flat plate behind the metanatal groove .

3- Pronotum and mesonotum were usually distinct.

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Plate (XIII): Phediole sinaitica the big headed ant ( Major worker)

Plate (XIV): Phediole sinitica (Minor worker)

Plate (XV): Phediole pallidula

Plate (XVI): Phediole indica

4.1.4.2. Pheidole sinaitica Mayr (1862) In this species the postpetiole was about as wide as long. The second segment of the funiclus was slightly longer than wide Plate (XIII). The whole body was reddish-brown in colour.

The workers of this species were found in two different forms a major worker, and a minor worker.

The Major worker of this species was twice as big as the minor worker. The major worker was found in only one locality namely in the farm near Elsemier village south west of Khartoum. Both minor and major workers were found in the same colony. This indicates that the major workers probably performed a defensive role in the colony Plate (XIV).

4.1.4.3. Pheidole pallidula Nylander (1849)

Head with quite deep occipital scalloping.

The lateral propodeum and petiole were smooth and shiny and the overall colour was pale yellowish-brown Plate (XV).

4.1.4.3. Pheidole indica Mayr 1879.

The head and gaster were dark in colour while the rest of the body was yellowish or reddish –brown. The thorax was anteriorly smooth and polished and a little narrower than the head and the abdomen was smooth and shiny Plate (XVI).

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Figure (8): Genus: Phediole

42

Plate (XVII): Tetramorium sericeventre

Plate (XVIII): Camponotus oasium

4.1.5.Genus: Tetramorium Fig (9)

4.1.5.1.Characteristics

1- The head was not heart-shaped in full face view Fig. (9)

2- The eyes were only rarely behind midlength of the sides of head Fig. (9).

3- The propodeum was usually armed with a pair of spines or teeth Fig (9).

4.1.5.2. Tetramorium sericeventre Emery 1877

Head, alitrunk and pedical densy and fine punctate with over rugoreticulation Plate (XVII). Gaster with extremely fine puncturation. Propodeal spines relatively long, narrow and triangular Plate (XVII).

The dorsal side of the propodeam was without hairs. The general colour was orange-brown, except the gaster which was black Plate (XVII).

4.2. Subfamily: Formicinae

In this subfamily four genera and eleven species were found.

4.2.1. Genus: Camponotus Fig (10)

4.2.1.1.Characteristics

Camponotus is a polymorphic genus, which is characterized by the following:

1- The petiole was a node or a scale which was never armed with teeth or spines.

2- The propodeum was unarmed.

3- The gaster was not capable of reflexion over the alitrunk.

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Plate (XIX): Camponotus (Tanaemyrmex) maculatus

Plate (XX): Camponotus (Orthonomyrmex) sericeus

Plate (XXI): Camponotus (Myrmopiromis) niveoetosus

Plate (XXII): Camponotus species 5

4- The mandible was usually with seven teeth at most, sometimes fewer and only very rarely with more than seven teeth .

4.2.1.2. Camponotus oasium Forel (1890)

The petiole of the species is rounded Plate (XVIII).

4.2.1.3.Camponotus (Tanaemyrmex) maculatus Fabricius (1781).

Yellow transverse patches on gaster appear as a row of two or three pale spots; subcephalic hairs numerous Plate (XIX).

4.2.1.4. Camponotus (Orthonomyrmex)sericeus Fabricius (1798).

Whole body hairy and propodeum broadly dentate Plate (XX).

This species was always found feeding on honey dew and was always associated with plants infested with aphids, scale insects and mealy bugs.

4.2.1.5. Camponotus (Myrmopiromis) niveoetosus Mayr (1862).

Gaster has very fine and dense striations, covering of hairs being strong but blunt. The dorsal pronotum is clearly bordered on the anterior edges Plate (XXI).

4.2.1.6. Camponotus species 5. Plate (XXII).

The whole body dark, the species in medium size, gaster with golden or white hairs, found as foragers on the ground, common in Khartoum University.

4.2.1.7. Camponotus species 6. Plate (XXIII).

The thorax of this species was thin and this species have smaller size than five species of camponotus recorded in this study.

The head and gaster were dark and thorax and petiole were brown in colour.

44

Figure (9): Genus: tetramorium

Figure (10): Genus: Camponotus

45

Plate (XXIII): Camponotus species 6

Plate (XXIV): Cataglyphis desertorum (sun dog)

Plate (XXIII): Camponotus species 6

Plate (XXIV): Cataglyphis desertorum (sun dog)

4.2.2. Genus: Cataglyphis Fig (11)

4.2.2.1. Characteristics

1- The metapleura have a distinct wide orifice for the metapleural gland, situated above the hind coaxa and blow the level of the propodeal spiracle.

2- Propodeal spiracle an elongate vertical or near- vertical slit Plate.

3- Ocelli present see .

4- First gasteral sternite in ventral view with a transverse sulcus immediately behind the helcium .

Members of this genus seemed to prefer dry desert and semi desert localities and were usually found moving on the surface of the ground even in the hot mid-day when the ground temperature was very high. The local name for members of this genus (Kalb Alhar = sun dog) possibly refers to this habit.

4.2.2.2. Cataglyphis desertorum Forel 1894

The workers had short legs and were very dark red in colour. They have well-developed spines on the tibiae Plate (XXIV).

4.2.2.3. Cataglphis lividus Andre (1881)

The whole body was yellow, and this species was smaller than Cataglyphis desertorum Plate (XXV).

4.2.3. Genus: Lepisiota Fig (12)

4.2.3.1. Characteristics

1- Antennae were formed of eleven segments each Plate (XXVI).

46

Plate (XXV): Cataglyphis lividus

Plate (XXVI): Lepisiota gracilicornis

Plate (XXVII): Lepisiota semenovi

Plate (XXVIII): Paratrechina longicornis

2- Propodeum was armed with a pair of spines, a pair of teeth, or a pair of tubercles.

3- The dorsal edge of petiole was usually armed with a pair of teeth or a pair of spines .

4.2.3.2. Lepisiota gracilicornis Forel 1892

The propodeal spines were short. Alitrunk was long thin and the antennal scapes were twice as long as the head width Plate (XXVI).

4.2.3.3. Lepisiota semenovi Ruzsky (1905)

The head was squire in shape with sparse erect hairs. The gaster was black or dark brown in colour. The alitrunk and also most of legs were red or reddish-yellow in colour Plate (XXVII).

4.2.4. Genus: Paratrechina Fig (13)

4.2.4.1. Characteristics

1- Ocelli were absent Fig. (13) and Plate (XXVIII).

2- First gastral sternite immediately behind the helicum entire, without a transverse sulcus .

3- Eyes large and very conspicuous Plate (XXVIII) and Fig. (13).

4.2.4.2. Paratrechina longicornis Latreille 1802

The antennal scapes were very long, extending as far as the mesonotum. The general body colour was brown to black. The eyes were large and the legs were long. The petiole was low Plate (XXVIII).

47

Figure (11): Genus: Cataglyphis

Figure (12): Genus : Lepisiota

48

Figure (13): Genus : Paratrechina

49

Plate (XXIX): Anochetus traegaordhi

Plate (XXX): Pachycondyla sennaarensis

4.3. Subfamily: Ponerinae

Two genera were recorded from this primitive subfamily.

4.3.1. Genus: Anochetus Fig (14)

4.3.1.1. Characteristics

1- In this genus the nuchal carina formed a broad, uninterrupted curve across the posteroidorsal extremity of the head.

2- The median groove was usually absent or ill-defined and shallow Plate (XXIX) and Fig. (14).

3- Dorsal most tooth of apical mandible series was acute.

4.3.1.2. Anochetus traegaordhi Mayr 1904

The thorax was very smooth and the petiole tapered to a dorsal crest. The propodeum was long and the mandibles were inserted in the middle of the front margin of the head.

The general body colour was yellowish-orange, slightly darker on the mid gaster Plate (XXIX).

4.3.2. Genus: Pachycondyla Fig (15)

4.3.2.1. Characteristics

1- The mandible was armed with five or more teeth.

2- Helcium was located very low on the front of the first gastral segment so that the first gastral segment had a long vertical anterior face in profile.

3- The tibiae of middle and hind legs had each one large pectinate spur and another small, simple spur.

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4.3.2.2. Pachycondyla sennaarensis Mayr (1862)

The workers of this ant have a deep mesopropodeal furrow and their mandibles had a dorsolateral pit. The eyes were relatively large. The overall body colour was black and the appendages had a deep red- brown colour Plate (XXX). This species is famous for its painful sting especially the winged reproductive females, which were common during the early rainy season.

4.4. Karyotypes of Camponotus (Myrmopiromis) niveoetosus and Camponotus species 6 :

4.4.1.Camponotus (Myrmopiromis) niveoetosus

The chromosomes number was 2n = 21

Fig. (16a) and Fig. (16b).

4.4.2. Camponotus sp6

The chromosomes number was 2n= 25

Fig. (17 a) and Fig. (17 b).

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Figure (14): Genus: Anochetus

Figure (15): Genus : Pachycondyla

52

Figure (16 a): A spread metaphase plate. The number of chromosomes was 2n=21

Figure (16 b): An Ideogram of Somatic Chromosomes of Camponotus niveoetosus arranged in pairs

53

Figure (17 a): A spread metaphase plate. The number of chromosomes was 2n=25

Figure (17b): An Ideogram of Somatic Chromosomes of Camponotus species6 arranged in pairs.

54

CHAPTER FIVE

DISCUSSION

The subfamilies of ants recorded in Khartoum during this study were the Myrmicinae (five genera and twelve species) followed by the Formicinae (four genera and eleven species) and the Ponerinae (two genera and two species). In the present study eleven genera of ants were recorded in Khartoum two of which, Lipsiota and Paratrechina were not previously recorded. In previous studies fourteen genera of ants were found in Khartoum. Five of these genera namely: Messor, Cataulatus, Leptothorax, Dorylus, Aenictus were not found during this study. On the other hand twenty five species of ants were found during this study compared to twenty one species in previous records.

Dorlylus fimbriatus laevipodex has been found in Khartoum by Santschi (1933); in July 2005, during the raining season, one male specimen was collected at night from under an electric light. Males of this genus are winged and large in size, generally known as sausage ants. Members of this species are nomadic predatory ants which do not build permanent nests and are usually called army ants. No workers or queens of this species have been found and therefore this genus has not been included in the list of Khartoum ants.

Species of the small sized ants: Monomorium and Tetramorium were always associated with humans and were found in almost in every house garden and farm. They are considered very irritating pests and are usually fond of sweet food or food rich in protein such as meat. Their ability to look for and find food indicates that they could possibly pollute food with agents of human diseases specially intestinal parasitis. This has to be confirmed through further research.

55

Monomorium species are famous for their ability to form new colonies any wherever they can find a suitable cover such as under a carpet or under apiece of wood or a piece of cardboard. This is because about 20% of every established colony is formed of secondary reproductives or fertile females, which cannot be differentiated externally from ordinary workers.

Any small group of individuals can separate from the original colony and the few reproductives found in it can lay eggs which hatch into infertile workers and as a result a new colony is established.

Pachycondyla sennaarensis was also always associated with human dwellings and gardens and they are very well known because of their painful stings especially those delivereded by winged reproductive females when the swarm at the beginning of the rainy season. This species needs further work to investigate the long-lasting painful venom they inject in people.

Species like Camponotus maculatus (or sugar ant) may have some importance because they may transfer disease through their feeding on sweet food.

Camponotus sericus can be of importance to farmers and gardeners because it has been noticed taking honeydew from Aphids nerri as well as mealy bugs Cloudesly –Thompson (1964). Ants which feed on honeydew protect the species that produce it and transport them from one plant to another to keep a continuous supply.

The karyotyping studies of two species of genus Camponotus agree with Crozier 1975 he said the known chromosome numbers of genus camponotus range between 2n=18 and 50, and also agree with karyotype

56

of Indian camponotus which observed 2n= 20,24,26,32,34,35,39,40 and 44 (Imai et al 1984).

Thus it may be concluded that further studies on taxonomy, ecology, behavior, and economic importance of ants in Khartoum and in Sudan in general is needed.

57

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Appendix 1: Letters Indicating Collection sites of ants in Khartoum State.

No. LOCALITY CODE

1 Blue Nile southern River Bank A

2 Khartoum Aciacia Forest B

3 Botanical Garden C

4 Soba Farm D

5 Tuti Island E

6 Khartoum University Main Campus F

7 Kukuo Research Farm G

8 Shambat H

9 Rawakeeb Research Station I

10 Salha Village J

11 Umbada - Omdurman K

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Appendix 2: Data about collection sites and species collected in Khartoum State.

species locality latitude longitude elevatio localit collection date microhabit habitat collected n y code codes at by

Anochetus Sudan; 15°38.885′ 32°27.599′ 416 m k ako 8 25-5- ground nest human houses a.k. traegaordhi Khartoum state, 2004 (semi desert) omer Umbada(west Khartoum)

Pachycondyla sudan;khartoum 15°36.789′ 32°32.366′ 379 m a ako 3 27-4- low river bank a.k. sennaarensis state;river bank 2004 vegetation vegetation omer

Lepisiota semenovi sudan; 15°40. 336′ 32°31. 520′ 392 m h ako 13 17-7- ground university a.k. khartoum state; 2004 forager(s) farms omer shambat(north vegetables khartoum)

Cataglyphis sudan; 15°22.631′ 32°39.725′ 400 m d ako 9 26-5- ground farm in semi a.k. desertorum khartoum state; 2004 forager(s) desert area omer elsemier village (soba farm) (south khartoum)

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Crematogaster sudan; 15°35.895′ 32°29.880′ 380 m b ako 4 5.5. ground nest acacia nilotica a.k. mimosae khartoum state; 2004 forest reserve omer khartoum sunt forest Monomorium sudan; 15°35.895′ 32°29.880′ 380 m b ako 5 12.5. dead branch acacia nilotica a.k. bicolor khartoum state; 2004 above forest reserve omer khartoum sunt ground forest Paratrechina sudan; 15°36.643′ 32°32.500′ 402 m f ako 2 24.4. ground nest khartoum a.k. longricornis khartoum state; 2004 university omer khartoum main campus university main campus Llepisiota sudan; 15°33.375′ 32°27.127′ 410 m j ako 10 17.6. ground nest desert area a.k. grcilicornis khartoum state; 2004 omer alsalha( west khartoum) Camponotus sudan; 15° 40.336′ 32° 31.520′ 392 m h ako 11 23-6- low university a k omer (orthonotomyrmex) khartoum state; 2004 vegetation farms sericeus (north vegetables khartoum); shambat Camponotus sudan; 15° 40.336′ 32° 31.520′ 392 m h ako 11 23-6- low university a k omer (myrmopiromis) khartoum state; 2004 vegetation farms ذذniveosetosus (north vegetables khartoum); shambat

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Camponotus sp5 sudan; 15° 36.066′ 32° 30.572′ 396 m c ako 14 4-8- low trees and a k omer khartoum state, 2004 vegetation garden plants elmogran from all region (botonical of the sudan garden)

Monomorium sudan; 15° 36.643′ 32°32.500′ 402 m f ako 2 24-4- ground nest khartoum a k omer areniphilum khartoum state; 2004 university a k omer khatoum main campus university main campus Monomorium sudan; 15° 36.789′ 32° 32.366′ 379 m a ako 1 21-4- rotten log river bank a k omer destructor khartoum state; 2004 vegetatation river bank Monomorium sudan; 15° 40.336′ 32° 31. 520′ 392 m h ako11 23-6- low university a k omer salomonis khartoum state; 2004 vegetation farms shambat (north vegetables khartoum) Monomorium mayri sudan; 15° 37.144′ 32° 34.836′ 397 m g ako 13 17-7- rotten log mix farm a k omer khartoum state; 2004 (vegetation, kukoo;(east cattle, furite khartoum) and cattle farm) Tetramorium sudan; 15° 36.643′ 32° 32.500′ 402 m f ako 7 20-5- under tree khartoum a k omer sericeventre khartoum state: 2004 bark university khartoum main campus university main campus

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Pheidole pallidula sudan; 15° 40.336′ 32° 31.520´ 392 m h ako 11 23-6- low university of a k omer khartoum state; 2004 vegetation khartoum farm shambat; (north vegetables khartoum) Pheidole sinaitica sudan; 15° 22.631′ 32° 39.725′ 400 m d ako 9 26-5- ground nest farm in semi a k omer (major worker) khartoum state; 2004 desert area elsemier village (soba farm) ;( south khartoum) Pheidole indica sudan; 15° 37.144′ 32° 34.836′ 397 m g ako 13 17-7- rote pocket mix farm a k omer khartoum state; 2004 above (vegetables, kukoo; (east ground cattle, furite khrtoum) and cattle farm)

Pheidole sinaitica sudan; 15° 33.375′ 32° 27.127′ 410 m j ako 16 7-8- ground nest desert area a k omer (minor worker) khartoum state; 2004 salha (west khartoum) Cardiocondyla sudan; 15° 35.895′ 32°29.880′ 380 m b ako 5 12-5- rotting log acacia nilotica a k omer emeryi khartoum state; 2004 forest reserve khartoum sunt forest Cardiocondyla sudan; 15° 35.895′ 32° 29.880′ 380 m b ako 5 12-5- rotting log acacia nilotica a k omer elegans khartoum sunt 2004 forest reserve forest

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Camponotus sudan; 15° 36.643′ 32° 32.500′ 402 m f ako 7 20-5- ground khartoum a k omer oasium khartoum state; 2004 foragers university khartoum main campus university main campus

Cataglyphis lividus sudan; 15° 2′ 32° 0′ 420 m i ako 15 28-7- on tree desert area a k omer khartoum state; 15° 36′ 32° 10′ 2004 trunk rawakeeb (west khartum) 45 km from khartoum Camponotus sudan; 15° 36.789′ 32° 32.366′ 379 m a ako 3 27-4- ground river bank a k omer (tanaemyrmex) khartoum state; 2004 foragers maculatus river bank

Camponotus sp6 sudan; 15° 36.643′ 32° 32.500′ 402 m f ako 2 24-4- ground khartoum a k omer khartoum state; 2004 foragers+ university khartoum low main campus university main vegetation campus

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اﻟﺨﻼﺻـــــﺔ

ﺠﻤﻊ ﺨﻤﺴﺔ ﻭﻋﺸﺭﻴﻥ ﻨ ﻭ ﻋ ﺎﹰ ﻤﻥ ﺍﻟﻨﻤل ﻤـﻥ ﻤﻨـﺎﻁﻕ ﻤﺨﺘﻠﻔـﺔ ﻓـ ﻲ ﻭﻻﻴـﺔ ﺍﻟﺨﺭﻁﻭﻡ. ﺼﻨﻔﺕ ﻫﺫﻩ ﺍﻻﻨﻭﺍﻉ ﺇﻟﻰ ﺜﻼﺙ ﺘﺤﺕ ﻋﻭﺍﺌل ﻭﺍﺤﺩﻱ ﻋﺸﺭ ﺠ ﻨ ﺴ ﺎﹰ. ﻋﺭﻓﺕ ﺜﻼﺜﺔ ﻭﻋﺸﺭﻴﻥ ﻤﻥ ﺍﻟﺨﻤﺴﺔ ﻭﺍﻟﻌﺸﺭﻴﻥ ﺇﻟﻰ ﻤﺴﺘﻭﻱ ﺍﻟﻨﻭﻉ، ﻫﻨﺎﻟـﻙ ﺠﻨـﺴﺎﻥ ﻴﻀﻤﺎﻥ ﺜﻼﺜﺔ ﺃﻨﻭﺍﻉ ﻭﺠﺩﺕ ﻟﻠﻤﺭﺓ ﺍﻻﻭﻟﻰ ﻓﻲ ﺍﻟﺨﺭﻁﻭﻡ. ﺇﻋﺘﻤﺩ ﺘﻌﺭﻴﻑ ﺍﻟﻌﻴﻨﺎﺕ ﻟﻠﺠﻨﺱ ﻭﺍﻟﻨﻭﻉ ﻋﻠﻰ ﺍﻟﻭﺼﻑ ﺍﻟﺨﺎﺭﺠﻲ ﺘﻡ ﺍﻟﺘﻌـﺭﻑ ﺒﻤﺴﺎﻋﺩﺓ ﻤ ﺨﺘﺼﻴﻥ ﻓﻲ ﻋﻠﻡ ﺘﺼﻨﻴﻑ ﺍﻟﻨﻤل ﻓﻲ ﻜل ﻤﻥ ﺒﺭﻴﻁﺎﻨﻴﺎ ﻭﺍﻟﻭﻻﻴﺎﺕ ﺍﻟﻤﺘﺤـﺩﺓ ﺍﻻﻤﺭﻴﻜﻴﺔ.

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