(Heteroptera: Nepomorpha: Naucoridae) of Tanzania

TAXONOMIC COMPOSITION AND DISTRIBUTION OF SAUCER BUGS

(HETEROPTERA: NEPOMORPHA: NAUCORIDAE) OF TANZANIA

A Thesis Presented to The Faculty of the Graduate School At the University of Missouri

In Partial Fulfillment Of the Requirements for the Degree Master of Science

By AARON YESAYA MBOGHO Dr. Robert W. Sites, Thesis Supervisor

DECEMBER 2012

The undersigned, appointed by the dean of the Graduate School, have examined the Thesis entitled TAXONOMIC COMPOSITION AND DISTRIBUTION OF SAUCER BUGS (HETEROPTERA: NEPOMORPHA: NAUCORIDAE) OF TANZANIA Presented by AARON YESAYA MBOGHO A candidate for the degree of Master of Science And hereby certify that, in their opinion, it is worthy of acceptance.

Dr. Robert W. Sites, Thesis Supervisor, Plant Sciences Entomology Area

Dr. Richard M. Houseman, Plant Sciences Entomology Area

Dr. Barry C. Poulton, Columbia Environmental Research Center

DEDICATION

……..to my late grandmother; Martha Hongoa, who laid the foundation for my education ACKNOWLEDGEMENTS

I am gratified to thank Professor Robert Sites, my advisor and thesis supervisor for his assiduous moral support and technical advice that equally made my studies at the

University of Missouri (MU) a success. His participation and practical guidance during insect collection trip in Tanzania and processing that led to description of two new species gave this work its contemporary shape. My heartfelt thanks extend to my committee members Dr. Richard Houseman and Dr. Barry Poulton for their support and advice during the entire period of my study that enabled me to successfully fulfill the requirements for the desired degree. My sincere gratitude equally extends to my employer; the Ministry of Agriculture, Food and Cooperatives, under the government of the United Republic of Tanzania for granting me a study-leave. It is this munificent leave of absence that made my on campus program of study in a distant continent achievable.

I would like to express my sincere thanks to all those who in one way or another shaped this work including my family members and friends. It is the generous Division of Plant Sciences assistantship and the Thurgood Marshall Fellowship, a prestigious academic award of the Graduate School that financially pillared me during the entire period of my studies in the United States. This work in particular was supported by

Freeda Thomas Yeo Agriculture Endowment Fund and Enns Entomology Museum of the

MU. Many individuals at the University of Missouri and the Tanzanian Ministry cooperated to make my study achievable and are hereby acknowledged to conclude the endless list of role players in this academic journey.

TABLE OF CONTENTS

ACKNOWLEDGEMENTS ...... ii

LIST OF TABLES ...... viii

LIST OF FIGURES ...... ix

ABSTRACT ...... x

CHAPTER 1 INTRODUCTION ...... 1

CHAPTER 2 LITERATURE REVIEW ...... 4

General history of research on Hemiptera and aquatic insects in Africa ...... 4

Overview of ecology of Naucoridae ...... 4

Overview of taxonomy of Naucoridae ...... 6

Research on Naucoridae in Africa ...... 6

General topography, hydrology, and climate in Africa ...... 9

Topography, hydrology, and climate in Tanzania ...... 11

CHAPTER 3 ANNOTATED LIST OF NAUCORIDAE ...... 13

Introduction ...... 13

Materials and Methods ...... 14

Results ...... 16

Annotated list of Naucoridae of Tanzania ...... 18

Subfamily Laccocorinae ...... 18

Aneurocoris insolitus Montandon ...... 18

Discussion ...... 18

Diagnosis ...... 19

iii

Distribution ...... 19

Material Examined ...... 20

Ctenipocoris africanus Poisson ...... 20

Discussion ...... 20

Diagnosis ...... 21

Distribution ...... 21

Material Examined ...... 21

Laccocoris limicola (Stål) ...... 21

Discussion ...... 22

Diagnosis ...... 22

Distribution ...... 22

Material Examined ...... 22

Laccocoris limigenus Stål ...... 23

Discussion ...... 23

Diagnosis ...... 23

Distribution ...... 24

Material Examined ...... 24

Laccocoris spurcus congoensis Poisson ...... 26

Discussion ...... 26

Diagnosis ...... 27

Distribution ...... 27

Material Examined ...... 27

Subfamily Naucorinae ...... 28 iv

Macrocoris flavicollis Signoret ...... 28

Discussion ...... 28

Diagnosis ...... 29

Distribution ...... 29

Material Examined ...... 30

Macrocoris sp...... 33

Discussion ...... 33

Diagnosis ...... 34

Distribution ...... 34

Material Examined ...... 34

Naucoris obscuratus Montandon ...... 34

Discussion ...... 34

Diagnosis ...... 35

Distribution ...... 35

Material Examined ...... 36

Neomacrocoris bondelaufa Sites ...... 36

Discussion ...... 37

Diagnosis ...... 37

Distribution ...... 38

Type Material Examined ...... 38

Additional Material Examined ...... 38

Neomacrocoris handlirschi (Montandon)...... 39

Discussion ...... 40 v

Diagnosis ...... 41

Distribution ...... 41

Material Examined ...... 41

Neomacrocoris karimii Poisson ...... 42

Discussion ...... 42

Diagnosis ...... 43

Distribution ...... 43

Material Examined ...... 43

Neomacrocoris parviceps (Montandon) ...... 43

Discussion ...... 43

Diagnosis ...... 44

Distribution ...... 44

Material Examined ...... 45

Neomacrocoris usambaricus Montandon ...... 46

Discussion ...... 47

Diagnosis ...... 47

Material Examined ...... 47

Neomacrocoris vuga Sites ...... 47

Discussion ...... 48

Diagnosis ...... 48

Distribution ...... 48

Type Material Examined ...... 49

Additional Material Examined ...... 49 vi

Distribution Overview ...... 49

Discussion ...... 50

CHAPTER 4 TAXONOMIC KEY ...... 54

Key to the species of Naucoridae of Tanzania ...... 54

BIBLIOGRAPHY ...... 63

APPENDIX ...... 70

VITA ...... 81

vii

LIST OF TABLES

1. Taxa of Naucoridae recorded from Africa exclusive of Madagascar (La Rivers, 1971, 1974, 1976; Sites and Mbogho 2012) ...... 7

2. Checklist of Naucoridae of Tanzania ...... 17

3. Taxa of Naucoridae found in Tanzanian Land Resource Zones. Species not collected in this study but known from Tanzania are marked with an asterisk ...... 50

viii

LIST OF FIGURES

1. Land resource zones map of Tanzania showing collection localities of aquatic insect sampling. LV, Lake Victoria; LT, Lake Tanganyika; LR, Lake Rukwa; LN, Lake Nyasa ...... 16

2–3. Head showing the difference between subfamilies. (2) Laccocorinae (Laccocoris limigenus) with front folded posteroventrally, (3) Naucorinae (Neomacrocoris parviceps) not folded ...... 58

4–6. (4) Aneurocoris insolitus, (5) Ctenipocoris africanus, (6) Naucoris obscuratus ...... 58

7–9. Species of Laccocoris. (7) Laccocoris limicola, (8) Laccocoris limigenus, (9) Laccocoris spurcus congoensis ...... 59

10–16. Species of Neomacrocoris. (10) Neomacrocoris bondelaufa, (11) Neomacrocoris handlirschi, (12) Neomacrocoris karimii, (13) Neomacrocoris parviceps ocellatus, (14) Neomacrocoris parviceps parviceps, (15) Neomacrocoris usambaricus, (16) Neomacrocoris vuga ...... 60

17–18. Species of Macrocoris. (17) Macrocoris flavicollis, (18) Macrocoris sp. (undescribed) ...... 61

19–20. Female subgenital plates. (19) Neomacrocoris usambaricus, (20) Neomacrocoris vuga ...... 61

21–24. Male genitalia. (21) Neomacrocoris bondelaufa, (22) Neomacrocoris parviceps, (23) Neomacrocoris karimii, (24) Neomacrocoris vuga...... 62

25–26. Lateral view of pronotum. (25) Neomacrocoris handlirschi, (26) Neomacrocoris parviceps ...... 62

ABSTRACT

Naucorids are predaceous aquatic insects and the principal arthropod predators of biting fly larvae. Naucorids exert natural control of mosquito larvae and are important with respect to malaria in Tanzania. However, naucorids have been documented to harbor the mycobacterium responsible for Buruli ulcer in Africa, although transmission is not known. Therefore, to provide a sound foundation for future biological, ecological, and epidemiological investigations, a study to document the taxonomic composition and distribution of the species of Naucoridae of Tanzania was conducted from 2010 to 2012; a total of 127 localities sampled. Habitats sampled included streams, rivers, waterfalls, ponds, lakes, reservoirs, swamps, and cold and hot springs. Previously, only four genera and seven species were known from Tanzania. All taxa previously recorded from

Tanzania were re-collected except two species (Neomacrocoris karimii Poisson and N. usambaricus Montandon). These collections added an additional two genera and eight species, including two undescribed species of Neomacrocoris and one undescribed species of Macrocoris. As a result, the known fauna of Naucoridae of Tanzania now comprises 6 genera and 14 species. This study provides the first authoritative identification resource to determine the entire fauna of Naucoridae of an African country using modern techniques. Publication of this study can serve as a model for future studies of this type in other countries in the African continent.

CHAPTER 1 INTRODUCTION

The natural world includes a myriad of life forms, many of which have direct effect on mankind in various ways. Some species of plants, animals, and fungi are used by humans for food, shelter, clothing, and energy; research is underway to assess the use of algae for oil production (Amin 2009). Other species act as competitors with man for the same resources. Still other species directly affect the health of humans by vectoring disease agents. In the absence of mankind, these species interact with one another in a more or less sustainable fashion. However, to proceed with human development in relative ignorance of the biota inhabiting a region carries with it the potential loss of useful taxa as well as the possible embellishment of those that might be detrimental. This says nothing of the cascade of effects with the ensuing trophic imbalance. Inventories of regional biota represent an important and frequently overlooked area of research because many scientists and others are more interested in focusing on the taxa of obvious human importance while ignoring those presumed to be inconsequential. However, those species thought to be unimportant could have profound effects that might not be readily apparent. As such, inventories are a necessary and important first step in understanding the biota of a region. Several inventories concerning aquatic Heteroptera have been conducted at high profile national parks and other areas in Africa, although many of these studies are old, incomplete, and of dubious accuracy. The family Naucoridae has not been inventoried in any region of Africa, except as a component of taxonomically broader surveys of aquatic Heteroptera, and never in Tanzania.

Naucorids are of contrasting economic importance because they are both beneficial and potentially harmful. Their beneficial aspect is in their feeding on the larvae of noxious flies. They have been reported as effective predators of mosquito larvae and pupae, including species of Anopheles (Wladimirow and Smirnov 1932), which vector malaria in Africa, hence suggesting them as possible biological control agents of malaria vectors. Conversely, they are considered to have potentially harmful effects on human health as they have been documented to carry the mycobacterium responsible for Buruli ulcer, a necrotizing “skin eating" disease (Merritt et al. 2010,

Ebong et al. 2012). In Africa, Buruli ulcer has received the greatest notoriety in the West

African countries of Ghana and Benin, although it occurs elsewhere with cases documented in Kenya and Uganda, just north of the Tanzanian border (Walsh et al.

2009).

Saucer bugs belong to family Naucoridae (Heteroptera) and are readily distinguished from other insects by the shape of the body; Naucorids are dorsoventrally flattened, ovate, and have retentorial prothoracic legs and natatorial meso- and metathoracic legs (Sites 2000). Naucorids range in length from 0.4 to 1.9 cm. Other distinctive features are that the membrane of the hemelytra is without veins and the abdomen does not have terminal filaments. In most heteropterans, the membrane contains veins, of which the number and arrangement serve to distinguish some families.

Naucorids pass all their life stages underwater and are predaceous, feeding on other animals that they can overpower. Most species of Naucoridae inhabit the tropics although a few are in temperate regions. They are predominant members of macroinvertebrate communities in tropical streams and are believed to greatly influence

2 the biota of the streams. However, the ecological parameters associated with these insects are not well known (Sites 2000). Little is known about the naucorid fauna of

Tanzania other than scattered distribution records (e.g., Poisson 1949). This study explored the species of Naucoridae and their distribution throughout the country including in Lake Tanganyika, which forms the western border.

The goal of this study was to conduct a faunistic survey of the species of Naucoridae of

Tanzania, identify the extant species, and document their distribution.

The specific objectives were: i. To determine the species composition of the naucorid fauna of Tanzania. ii. To document the distribution of each species of naucorid in Tanzania. iii. To prepare a taxonomic key for the identification of the naucorids Of Tanzania.

CHAPTER 2 LITERATURE REVIEW

General history of research on Hemiptera and aquatic insects in Africa

Our knowledge of the Hemiptera fauna of Africa, especially south of the Sahara, is poorly known (Linnavuori 1971). The extensive works on African Hemiptera are those of the 19th and 20th centuries by Stål: Hemiptera Africana (1864-1865) and Enumeratio

Hemipterorum (1870-1876) (Linnavuori 1971). So far, the best researched region in tropical Africa is Congo whose fauna was treated extensively by Schouteden in his numerous publications (Linnavuori 1971). In addition to Schouteden, Villiers studied the

Reduviidae of the former French colonies in Africa culminating with Hemipteres

Reduviides de l’ Afrique Noire in 1948. Mancini published several works on the fauna of

Ethiopia and other Italian colonies. In 1967, Villiers again revised the Reduviidae of

Africa in his paper Revision des Reduviidae Africains. Only scattered papers have been published from East Africa (Linnavuori 1971) including Hynes (1955) notes on East

African aquatic Heteroptera, although this suggests that there is much to be done, including in Tanzania. The South African fauna has been treated in various older papers on individual taxa [e.g., Notonectidae and Corixidae (Hutchinson 1929)] as well as in the contemporary South African Animal Life series, which includes the Field Guide to

Insects of South Africa (Picker et al., 2002).

Overview of ecology of Naucoridae

Naucorids are contrastingly both beneficial and potentially harmful. Their beneficial

4 aspect is in their feeding on the larvae of noxious flies. Naucorids are predaceous and pass all their life stages underwater feeding on other animals that they can overpower.

They are predominant members of macroinvertebrate communities in tropical streams and are believed to influence greatly the biota of the streams. However, the ecological parameters associated with these insects are not well known (Sites 2000).

Naucorids have been reported as effective predators of larvae and pupae of mosquitoes, including species of Anopheles (Wladimirow and Smirnov 1932), which vector malaria.

Conversely, they have potentially harmful effects on human health in that they have been documented to carry in their salivary glands the mycobacterium responsible for Buruli ulcer, a necrotizing “skin eating" disease (Merritt et al. 2010). In Africa, Buruli ulcer has received the greatest notoriety in the West African countries of Ghana and Benin, although it occurs elsewhere in Africa with cases documented in Kenya and Uganda, just north of the Tanzanian border (Walsh et al. 2009). After tuberculosis, Buruli ulcer is emerging as the second most frequently diagnosed mycobacterial disease in humans, and it remains a neglected disease even though it has been reported recently in some developed countries (Merritt et al. 2010). By 2010, more than 32 countries had reported

Buruli ulcer cases, most of which were in West Africa, Southeast Asia, Central and South

America, the western Pacific, Australia, and China. The disease manifests itself as large skin lesions that result in scarring and deformities. When severe infections are sustained, amputations and disabilities commonly result. The disease is most frequently reported in children from 4 to 15 years old, especially in Africa (Merritt et al. 2010). Mycobacterium ulcerans is more frequent in communities in the vicinity of rivers, swamps and wetlands.

The disease is thought to be associated with anthropogenic changes in the aquatic

5 environment, such as a result of environmental disturbance including deforestation, dam construction, and agriculture. This is considered a ‘‘mysterious disease’’, with its mode of transmission unclear (Merritt et al. 2010), although bites from aquatic Heteroptera, including Naucoridae, have been suggested as the mode of transmission (Ebong et al.

2012).

Overview of taxonomy of Naucoridae

The current taxonomic arrangement of 5 subfamilies with 37 genera and 378 species is the result of periodic additions of genera to a long-standing set of subfamilies. As many as 7 subfamilies have been recognized, although 2 of these recently have been elevated to family level (Aphelocheiridae and Potamocoridae). The classification of the family as of

1971 was presented as a catalog and was updated twice in subsequent years (La Rivers

1971, 1974, 1976). Štys and Jansson (1988) later provided information and clarification regarding the status of subfamily, tribe and generic taxonomy.

Research on Naucoridae in Africa

The naucorid fauna in Africa presently is known to include 56 species in 8 genera and 2 subfamilies. Little is known about the naucorid fauna of Tanzania other than scattered distribution records of only 7 species in 4 genera in both subfamilies (e.g., Signoret 1860;

Montandon 1897a, 1900, 1909, 1913a, b; Poisson 1949, 1963). Thus, the literature from which one can begin to understand the naucorid fauna in Tanzania is old and cursory at best.

Table 1. Taxa of Naucoridae recorded from Africa exclusive of Madagascar (La Rivers, 1971, 1974, 1976; Sites and Mbogho 2012).

Taxon Distribution1

Laccocorinae

Aneurocoris (Aneurocoris) insolitus Montandon TZ

Aneurocoris (Aneurocorisella) marlieri Poisson DRC

Ctenipocoris africanus Poisson CO

Heleocoris linnavuorii amaricus Linnavuori SU, ER

Heleocoris linnavuorii linnavourii Poisson SU, ET, ER

Laccocoris angustus Linnavuori MZ

Laccocoris bouvieri Poisson BN, NG

Laccocoris chinai Poisson ET, DRC, SA

Laccocoris discus Montandon CM

Laccocoris dissidens Montandon AF

Laccocoris limicola (Stål) SA

Laccocoris limigenus Stål AF

Laccocoris nymphaerum Rochebrune AF

Laccocoris omoensis Linnavuori ET

Laccocoris salina Poisson DRC

Laccocoris spurcus spurcus (Stål) SA, DRC

Laccocoris spurcus congoensis Poisson AF

Laccocoris tauorgae Sallier Dupin SO

Naucorinae

Macrocoris convexus Montandon CO, NG, SU, UG, SA

Macrocoris flavicollis chariensis Poisson CH

Macrocoris flavicollis flavicollis Signoret GH, RCA, BN, EA

Macrocoris flavicollis uelei Poisson CH, IV, SU

Taxon Distribution1

Macrocoris intermedius Poisson DRC

Macrocoris laticollis laticollis Montandon CM, AN

Macrocoris laticollis mongai Poisson CO

Macrocoris nigropunctatus africanus Poisson CA

Macrocoris n. nigropunctatus Montandon CA, NG, UV

Macrocoris schoutedeni Poisson CO

Macrocoris transvaalensis Distant SA

Naucoris ciliatistylus Linnavuori ET

Naucoris fuscipennis Schaum SEA

Naucoris kenyalis Poisson KE, ET, CO

Naucoris maculatus angustior Lethierry NA

Naucoris maculatus conspersus Stål NA

Naucoris maculatus maculatus Fabricius NA

Naucoris obscuratus nanus Poisson CO

Naucoris obscuratus obscuratus Montandon SU, ET, WA, EA, SA

Naucoris planus Germar AF

Neomacrocoris angusticeps (Haglund) CM

Neomacrocoris bondelaufa Sites TZ, SA

Neomacrocoris dubius Poisson WA

Neomacrocoris handlirschi (Montandon) DRC, MZ, RA, RCA, SA, SU, TZ, UG, ZM

Neomacrocoris katangae Poisson DRC

Neomacrocoris hungerfordi Poisson CO

Neomacrocoris karimii Poisson TZ

Neomacrocoris nanus Poisson DRC

Neomacrocoris ndugai Poisson RA

Neomacrocoris parviceps ocellatus Poisson CO, TZ

Taxon Distribution1

Neomacrocoris parviceps parviceps (Montandon) CO, DRC, GH, IV, SA, SU, TZ, UG, UV, ZI

Neomacrocoris poissoni Linnavuori IV, NI, SU

Neomacrocoris reavelli Sites SA

Neomacrocoris schaeferi Sites UG

Neomacrocoris usambaricus Montandon DRC, TZ

Neomacrocoris vaneyeni Poisson CM, DRC, IV

Neomacrocoris vuga Sites TZ

1AF, Africa; BN, Benin; CA, Central Africa; CH, Chad; CM, Cameroon; CO, Congo; DRC, Democratic Republic of

Congo; ET, Ethiopia; ER, Eritrea; GH, Ghana; IV, Ivory Coast; KE, Kenya; MZ, Mozambique; NA, North Africa; NI,

Nigeria; RA, Rwanda; RCA, Republic of Central Africa; SA, South Africa; SO, Somalia; SU, Sudan; TZ, Tanzania;

UG, Uganda; UV, Upper Volta; WA, West Africa; ZI, Zimbabwe; ZM, Zambia.

General topography, hydrology, and climate in Africa

The continent of Africa comprises 62 political territories with a total area of more than 30 million km2. The Mediterranean Sea separates Africa from Europe and the Red Sea separates much of Africa from Asia. High southern and eastern plateaus seldom fall below 600 m and have a mean elevation of approximately 1,000 m. The largest of these are the southern Great Karoo and Kalahari Desert. The South African plateau is connected to East Africa by distinct topographic features including mountain ranges, tablelands and depressions. The most striking features are two great lines of depression, due largely to the subsidence of whole segments of the Earth's crust, the lowest parts of which are occupied by vast lakes. Toward the south the two lines converge to one great valley occupied by Lake Nyasa. The Great Rift Valley is occupied for more than half its length by water: the Great Lakes of Tanganyika, Kivu, Edward and Albert. Within these 9 valleys are many volcanic peaks. The eastern depression is the East Africa Rift Valley with much smaller lakes, many of them brackish and without outlet. East of this rift valley are Mount Kilimanjaro (5,889m) and Mount Kenya (5,184 m).

The higher region of Africa is formed by the Ethiopian Highland, a mass of mountains forming the largest continuous area of high elevation in the continent (Fobil et al. 2003) with little of its surface below 1,500 m, whereas the summits reach heights of 4,600 to

4,900 m. This area includes the Ethiopian Rift Valley and lies northeast of the East

African Rift Valley, the northern continuation of which extends to the Red Sea. There are three main rivers in Africa: River Nile, the longest river on the continent; River

Congo, the highest water discharge on the continent; and River Niger, which flows out almost a half of its length through arid areas. The largest lakes are Lake Victoria and

Lake Chad in the center of the continent; Lake Tanganyika between the Democratic

Republic of Congo, Burundi, Tanzania and Zambia; and Lake Malawi stretching along the eastern border of Malawi. Numerous dams are situated throughout the continent, including Kariba in Zambia on Zambezi River, Asuan in Egypt on the River Nile, and

Akosombo, the biggest dam of the continent in Ghana on the Volta River (Fobil et al.

2003). Most of the continent is within the tropics and generally does not experience excessive temperature variations. However, arid regions including the lower plains and deserts experience greater temperature extremes; they can be much hotter and occasionally the temperature in the Sahara Desert can fall to the freezing point. In the extreme north and south the climate is warm temperate with the northern countries generally hotter and drier than those in the south. The southern part of the continent is narrower than the north and experiences a moderating influence of the surrounding

10 oceans. Within the equatorial zone, especially on the shores of the Gulf of Guinea and in the upper Nile basin, the continent experiences intensified rainfall.

The African continent generally south of the Sahara is the Afrotropical biogeographic realm, whereas northern Africa is part of the Palearctic biogeographic realm. Europe and much of Asia compose the rest of the Palearctic realm. It is only the Cape of Africa in

South Africa which is placed in the Afrotemperate region (Morrone 2002).

Topography, hydrology, and climate in Tanzania

Tanzania is in the eastern part of Africa, bordering the Indian Ocean between Kenya and

Mozambique. It lies between 6o 00’ and 35o 00’ E and has a total area of ca. 945,000 km2 with ca. 886,000 km2 of land and ca. 59,000 km2 of water bodies. The country has two offshore main islands of Zanzibar (Unguja and Pemba) and has 3,402 km of borders shared with eight countries: Burundi, 451 km; Kenya, 769 km; Malawi, 475 km;

Mozambique, 756 km; Rwanda, 217 km; Uganda, 396 km; Zambia, 338 km; and its coastline is 1,424 km. The climate in Tanzania varies from tropical along the coast to temperate in the highlands. The terrain is mainly plains along the coast and central plateau and highlands in north and south. The elevation extremes are the Indian Ocean and Mt. Kilimanjaro (5,895 m).

Land use is dominated by permanent pastures (40%) and forests and woodland (38%).

Only 3% of the land is arable, 1% is in permanent crops, and the remaining 18% represents other uses (Tanzania Bureau of Statistics 1993). Lake Tanganyika, Lake

Victoria, Lake Nyasa (Lake Malawi), and rivers such as Ruvu, Rufiji, Pangani, Mto-wa-

11 mbu plus natural and man-made dams and reservoirs are the main sources of irrigation and hydroelectric power.

CHAPTER 3 ANNOTATED LIST OF NAUCORIDAE

Introduction

The Hemiptera fauna of the world has been researched at different levels of thoroughness in different regions. For example, the faunas of Australia and North America are well documented. However, although the South African fauna has been treated (e.g., Picker et al. 2002), our knowledge of the Hemiptera fauna of most of Africa, especially south of the Sahara, is poorly known (Linnavuori, 1971). Much of the African continent is within the Afrotropical biogeographic realm and harbors a great diversity of tropical biota, with insects being the most diverse group. The Belgian Congo is the region of tropical Africa whose fauna has been most researched with explorations, which is reflected in several publications by various authors including Schouteden. Scattered minor papers have been published concerning the East African fauna (Linnavuori, 1971), suggesting there is much to be done in Africa, including in Tanzania. The naucorid fauna of Africa has not been systematically treated. The most recent works on the naucorid fauna of Tanzania date back to the colonial era and early 1960s when the Tanzanian mainland still was referred to as Tanganyika and the two Islands of Zanzibar referred to as Unguja and

Pemba. Unguja and Pemba joined to form Zanzibar, which in 1963 united with

Tanganyika to form the United Republic of Tanzania. Thus, Tanganyika, Unguja and

Pemba are valid place names prior to 1964, whereas Tanzania and Zanzibar are valid after 1964.

Materials and Methods

Extensive collecting was carried out in Tanzania for 25 consecutive days from 24 July to

17 August 2010, during which 121 sites were collected. Three additional sites: two in

Tanga Region on 28 December 2011 and one in Morogoro Region on 31 December 2011 were sampled; the same three sites were sampled again on 29 January and 1 February

2012 making a total of 127 collection sites in this study. Tanzania has been apportioned into six land resource zones (Panagos et al. 2011) and the collections were conducted in all six zones (Fig. 1). During all collections, naucorids were collected using aquatic D- nets (1.1 x 1.1 mm mesh) to sweep through submerged vegetation along shorelines and bank undercuts of various water bodies. Almost all accessible and safe water bodies encountered in the collection route were sampled. Sampled water bodies include rivers, streams, waterfalls, cold and hot springs, dams, reservoirs, lakes, ponds, marshes and pools. The collection localities were identified by L-numbers from 1144 to 1267, 1322 to

1324, and 1332 to 1334 (see details of locality data in Appendix). Photographs of the collection localities can be viewed in the Locality Image Database at the internet site of the Enns Entomology Museum, University of Missouri. One collection site, L-1155, was in Rwanda along the border with Tanzania, and another, L-1217, was a blacklight sample; thus, data for those two collections are excluded here. At each site, net contents were placed into a white plastic pan and all insects removed and placed into a jar containing 95% ethyl alcohol. Alcohol was changed later in the day to compensate for dilution. All samples were brought to the laboratory at the University of Missouri where naucorids were separated from other insects. Male genitalia were extracted under the microscope and placed into a genitalia vial containing glycerin. 14

The collecting route of travel in 2010 was mainly by road although air and water transportation also were used. The collection trip started in Mwanza Region on 24 July

2010 and finished in Kusini Region in Zanzibar on 17 August 2010. In Mwanza Region, collecting was conducted for two days on 24 and 25 July at 7 collection sites and in Mara

Region at 3 sites. On 26 and 27 July 2010, three sites sampled in Kagera Region and one site in Esta Province in Rwanda, then moved to Kigoma Region where collecting was conducted for 4 days from 27 to 30 July. Kigoma Region is bordered by Lake

Tanganyika. In Kigoma Region, a total of 26 collection sites including Lake Tanganyika and streams in Gombe National Park were sampled. From 30 July to 3 August, collecting was conducted in Rukwa Region and a total of 23 collection sites were sampled. Four collecting sites were sampled in Mbeya Region on 3 and 4 August, and four sites in

Ruvuma Region on 5 August. Two collecting sites in Iringa Region were sampled on 6

August, and seven collection sites were sampled in Morogoro Region on 6 and 7 August.

In Dodoma Region, two sites were sampled on 8 August. Six collection sites were sampled in Singida Region on 8 and 9 August. Four collection sites were sampled in

Manyara Region on 9 August. Six collection sites were sampled in Arusha Region on 10 and 11 August. Nine collection sites were sampled in Kilimanjaro Region from 11 to 13

August. Twelve collection sites were sampled in Tanga Region from 13 to 15 August; two collection sites were sampled in Pwani Region on 16 August. On 17 August, the team traveled by Sea Bus express boat to Zanzibar where four sites sampled in three different regions as follows: One site in Mjinimagharibi Region, two sites in Kaskazini

Region and one site in Kusini Region.

Figure 1. Land resource zones map of Tanzania showing collection localities. LV, Lake Victoria; LT, Lake Tanganyika; LR, Lake Rukwa; LN, Lake Nyasa (after Panagos et al. 2011).

Results

In this study, two subfamilies of Naucoridae were found: Laccocorinae and Naucorinae.

Collections of Laccocorinae comprised three genera: Aneurocoris, Ctenipocoris, and

Laccocoris and a total of 5 species. Collections of Naucorinae comprised three genera:

Macrocoris, Naucoris, and Neomacrocoris and a total of 7 species. Previously, the available literature indicated that only 4 genera and 7 species were known from Tanzania.

All taxa previously recorded from Tanzania were re-collected during data collection, with the exception of old records of two species from the Usambara Mountains. As a result of this study, two more genera, Ctenipocoris and Naucoris, and 5 more species are added to the naucorid fauna of Tanzania. Thus the known naucorid fauna now includes 6 genera and 14 species. Two of the species added to the fauna were undescribed in the genus

Neomacrocoris with distributions that are quite different; Neomacrocoris vuga Sites was collected in only one stream in the Usambara Mountains, whereas Neomacrocoris bondelaufa Sites was collected throughout western Tanzania in the East African Rift

Valley, although one specimen from South Africa also was examined. In addition, one undescribed species of Macrocoris was collected in Kagera Region.

In this chapter, a checklist and annotated list of species and subspecies of Naucoridae collected during this study and known from the literature are presented.

Table 2. Checklist of Naucoridae of Tanzania

Taxon Previous Record New Record

1. Aneurocoris insolitus Montandon x

2. Ctenipocoris africanus Poisson x

3. Laccocoris limicola (Stål) x

4. Laccocoris limigenus Stål x

5. Laccocoris spurcus congoensis Poisson x

Taxon Previous Record New Record

6. Macrocoris flavicollis Signoret x

7. Macrocoris sp. x

8. Naucoris obscuratus Montandon x

9. Neomacrocoris bondelaufa Sites x

10. Neomacrocoris handlirschi (Montandon) x

11. Neomacrocoris karimii Poisson x

12. Neomacrocoris parviceps (Montandon) x

13. Neomacrocoris usambaricus Montandon x

14. Neomacrocoris vuga Sites x

Annotated list of Naucoridae of Tanzania

Subfamily Laccocorinae

Aneurocoris insolitus Montandon

(Fig. 4)

Aneurocoris insolitus Montandon 1897a: Ver. Zool. Bot. Gesel. Wein, 47: 437.

Discussion

The genus Aneurocoris includes two species endemic to Lake Tanganyika: Aneurocoris insolitus Montandon and Aneurocoris marlieri Poisson. Label data of museum specimens indicated that A. insolitus was collected from rocks in this lake but during this collection it was not found among rocks. Irvine and Donohue (1999) indicated that this species lives on the bottom and on rocks, especially in shallow water, and possibly

18 remains submerged throughout its life cycle. This study found the reported habitat information to be inaccurate. Rather than living on or among rocks, this species was associated consistently with root mats of reed grass (Phragmites mauritianus Kunth) in the four sites where it was collected. Other sites with sand and rock substrates also were sampled, but A. insolitus was not found. The species was collected from the two extreme ends of the Tanzanian shoreline and one site in the middle. Polhemus & Polhemus

(2008) were in error in reporting that two endemic genera of Naucoridae are known from

Lake Tanganyika. Only one endemic genus (Aneurocoris) is known from this lake.

Poisson (1960c) provided data on A. insolitus and described a second species,

Aneurocoris marlieri, endemic to Lake Tanganyika from the northwestern shoreline in the Democratic Republic of Congo.

Diagnosis

Aneurocoris insolitus can be distinguished from other species of Naucoridae in Tanzania by the entirely coriaceous hemelytra which are devoid of veins and sutures. It is relatively small for the family, measuring 6.0–7.4 mm.

Distribution

Aneurocoris insolitus is limited to Lake Tanganyika, from Kigoma to the southernmost point in Tanzania. Streams and rivers which flow to the lake were not found to harbor the species. Its distribution in the lake is not continuous, but apparently restricted to the isolated patches of reed grass, Phragmites mauritianus, where the insects live among the rhizomes and root mats. The type specimen was listed from Mpala, Tanzania, which is not along the coastline of Lake Tanganyika. This is considered to be either a case of

19 mislabeling or perhaps another small village on the lakeshore by the same name existed at that time.

Material Examined

TANZANIA: KIGOMA Region: Lake Tanganyika, Kigoma, 04º 54.719'S 29º 37.989'E,

772 m, 28 July 2010, colls R. W. Sites & A. Mbogho, rootmats of matete on waveswept sandy shoreline, L-1166; same locality, 04º 55.319'S 29º 40.256'E, 773 m, 28 July 2010, colls R. W. Sites & A. Mbogho, dense rootmats of matete in heavy wave action, L-

1167. RUKWA Region, Lake Tanganyika, Karema, 06º 49.694'S 30º 26.646'E, 780 m,

31 July 2010, colls R. W. Sites & A. Mbogho, heavily wave swept roots of reed grass, L-

1189; Lake Tanganyika, Kasanga, 08º 26.746'S 31º 08.642'E, 773 m, 2 August 2010, colls R. W. Sites & A. Mbogho, waveswept reed grass clumps, L-1203.

Ctenipocoris africanus Poisson

(Fig. 5)

Ctenipocoris africana Poisson 1948: Rev. Zool. Bot. Afr., 41: 219–221.

Discussion

Ctenipocoris africanus is the only species of the genus known from Africa, but has not been reported from Tanzania previously. This species was described from Ituri in the

Democratic Republic of Congo (Poisson 1948). La Rivers (1960) subsequently redescribed it as Heleocoris faradjensis from Belgium Congo (=DRC), which later was synonymized with C. africanus (Polhemus and Polhemus 2008). This species is not common in collections but specimens from Uganda in the British Museum were examined. Presented here is the first record of C. africanus from Tanzania. 20

Diagnosis

This moderately-sized species measures 8.3 by 5.5 mm and can be recognized by heavily spined meso- and metatibiae and femora, and the profemora are relatively small. The greatest width of the pronotum is approximately twice its length at the midline, whereas

Aneurocoris and species of Laccocoris are three times as wide as long. The prothoracic tarsi have two segments in both sexes, whereas Aneurocoris and Laccocoris are sexually dimorphic; the male has two tarsal segments and the female only one. It is 8–9 mm in length.

Distribution

Ctenipocoris africanus is known only from Democratic Republic of Congo, Uganda, and

Tanzania.

Material Examined

TANZANIA: RUKWA Region: 110 km N of Mpanda, 05º 37.745'S 30º 36.039'E, 1295 m, 30 July 2010, colls R. W. Sites & A. Mbogho, shallow creek with muddy bottom and marginal grasses, L-1185. UGANDA: Namanve Swamp, Kampala, 18 i 1933, G.L.R.

Hancock, B.M. 1933-585, NA 10, 1-XI-32 / Pres. By Imp. Inst. Ent. B.M. 1933-585

(BMNH, 1♂); Kampala, 30 iii 1932, G.L.R. Hancock / Pres. By Imp. Inst. Ent. B.M.

1933-585 (BMNH, 1♀).

Laccocoris limicola (Stål)

(Fig. 7)

Naucoris limicola Stål, 1855: Ofv. Kong. Vet. Ak. Forh., 12: 46.

Laccocoris limicola: Stål, 1865, Holmiae, p. 178. 21

Discussion

This species was transferred from the genus Naucoris to Laccocoris by Stål (1865). The

African species of Laccocoris have been apportioned into two groups by Poisson (1949): the spurcus group and the limigenus group. Laccocoris limicola is in the spurcus group, which is characterized as having slightly shortened membranes of the hemelytra and subbrachypterous, brachypterous, or micropterous hindwings, and the posterolateral angles of the pronotum are more or less acute and recurved caudad (Poisson 1949).

Linnavuori (1987) added that the membranes of the hemelytra are acuminate apically.

Diagnosis

Laccocoris limicola can be distinguished from its two known congeners in Tanzania by the sharp posterolateral angles of the pronotum. The body length is 8.5–9.3 mm.

Distribution

Laccocoris limicola has been recorded from Gariep (Northern Cape Province) and

Limpopo, South Africa (Stål 1855). During these collections, the species was found in

Kagera and Kigoma regions of western Tanzania. In each of the collections, it was taken syntopically with L. limigenus Stål in association with rocks and submerged vegetation in moderate current.

Material Examined

TANZANIA: KAGERA Region; Kagera/Kigoma regional border, Muvozi River,

03º11.331'S 31º02.129'E, 27 July 2010, 1156 m, colls: R. W. Sites & A. Mbogho , rocky stream with dense filamentous vegetation, L-1158. KIGOMA Region; 40 km N of

Kigoma, 04º53.201'S 29º52.190'E, 952 m, 27 July 2010, colls: R. W. Sites & A. Mbogho, 22 rapid stream w/ slab rock gravel, marginal vegetation, L-1165.

Laccocoris limigenus Stål

(Fig. 8)

Laccocoris limigenus Stål, 1865: Holmiae, p. 178.

Laccocoris aurivillii Montandon, 1897b: Ann. Soc. Entomol. Belg., 47: 50–52.

Discussion

Laccocoris limigenus is arguably the most widespread and common species of

Laccocoris in Africa. L. aurivillii Montandon was considered by Poisson (1949) to be a race of L. limigenus, differing from the latter by having a larger body size and more slender phallosoma. Because of high variability in both characters, Linnavuori (1987) formally synonymized L. aurivillii with L. limigenus. Poisson (1949) placed L. limigenus in the limigenus group, which is characterized as being macropterous with alary dimorphism rare, and with the posterolateral angles of the pronotum rounded. Linnavuori

(1987) added that the membranes of the hemelytra are roundedly truncate. The species is common in oligotrophic rivers with a sandy bottom (Linnavuori 1981) and has been recorded in shallow pools in river sandbanks with dead leaves and detritus (Weir 1966).

The species was most frequently found in shallow streams with sandy or silty bottoms and in association with vegetation. It was collected syntopically with L. limicola at L-

1158 and L-1165.

Diagnosis

Laccocoris limigenus is distinguished easily from known congeners in Tanzania by the rounded posterolateral corners of the pronotum. Body length is variable and was reported 23 as 7.0–9.5 mm (Linnavuori 1987).

Distribution

This species occurs throughout Africa and is known specifically from Algeria, Ethiopia,

(Linnavuori 1971), Sudan (Linnavuori 1987), Chad, Nigeria (Linnavuori 1981) and Niger

(Poisson 1950), west to Benin (Poisson 1951, Linnavuori 1981) and Togo (Poisson

1951), south through Central African Republic (Linnavuori 1981), Kenya (Poisson 1963),

Democratic Republic of Congo (Poisson 1949, 1951, 1954; Linnavuori 1987), Tanzania

(Poisson 1960b, 1963), Zambia (Poisson 1963, Linnavuori 1987), Mozambique (Poisson

1934), and South Africa (Poisson 1963, Linnavuori 1987).

In Tanzania, it has been recorded from Arusha Region (as Maji ya Chai), Tanga

Region, and Usambara North (Poisson 1963). This species was found to be widespread and collected from 21 localities in 10 regions: Dodoma, Iringa, Kagera, Kigoma,

Manyara, Morogoro, Mwanza, Rukwa, Singida and Tanga regions.

Material Examined

TANZANIA: DODOMA Region: Lukali hot spring, 05º43.936'S 35º30.082'E, 991 m, 8

August 2010, colls: R. W. Sites & A. Mbogho, warm spring with algae and Typha angustifolia, L-1223. IRINGA Region: Silunga River at Chikuyu, 05º52.223'S

35º04.568'E, 842 m, 8 August 2010, colls: R. W. Sites & A. Mbogho, dead twigs in muddy river, L-1224; Mwiriwa River at Kimamba, 07º39.657'S 36º05.275'E, 1312 m, 6

August 2010, colls R. W. Sites & A. Mbogho, pooled stream w/ algae and overhanging grasses, L-1215. KAGERA Region: Kagera/Kigoma border, Muvozi River, 03º11.331'S

31º02.129'E, 1156 m, 27 July 2010, colls R. W. Sites & A. Mbogho, rocky stream with

24 dense filamentous vegetation, L-1158. KIGOMA Region: 40 km N of Kigoma,

04º53.201'S 29º52.190'E, 952 m, 27 July 2010, colls R. W. Sites & A. Mbogho, rapid stream w/ slab rock, gravel, marginal vegetation, L-1165. MANYARA Region: Yayeda

River at Bagheti, 04º00.158'S 35º20.470'E, 1974 m, 9 August 2010, colls R. W. Sites &

A. Mbogho, marginal overhanging grasses, L-1232; Tiawi, Kwaanseri Lake, 03º54.600'S

35º27.739'E, 1940 m, 9 August 2010, colls R. W. Sites & A. Mbogho, wetland with heavy vegetation, L-1234; MOROGORO Region: Ruaha River at Mbuyuni,

07º27.922'S 36º30.973'E, 527 m, 6 August 2010, colls R. W. Sites & A. Mbogho, large river w/ sparse marginal vegetation, L-1216; Magole River at Dumila, 06º23.221'S

37º21.629'E, 436 m, 7 August 2010, colls R. W. Sites & A. Mbogho, marginal veg in shallow sandy river, L-1222. MWANZA Region: Kisesa, ca. 20 km NE Mwanza,

02º33.266'S 33º03.767'E, 1222 m, 24 July 2010, colls, R. W. Sites & A. Mbogho, vegetation in pooled cattle stream, L-1145. RUKWA Region: 57 km N of Mpanda,

Masito/Ogala, 05º56.742'S 30º57.812'E, 1144 m, 30 July 2010, colls R. W. Sites & A.

Mbogho, slow stream, coarse roots at margin, L-1187; 1 km E of Karema, 782 m,

06º48.473'S 30º26.315'E, 31 July 2010, colls R. W. Sites & A. Mbogho, algal mat at margin, L-1190; Mpanda, Kasimba River, 06º21.602'S 31º04.382'E, 1064 m, 1 August

2010, colls R. W. Sites & A. Mbogho, grasses hanging into river, L-1195; Kamawe River at Puwi, 08º21.029'S 31º50.054'E, 1598 m, 3 August 2010, colls R. W. Sites & A.

Mbogho, clear stream, marginal vegetation, some rocks, L-1205. SINGIDA Region:

Kipumbuiko River, 05º05.667'S 34º45.940'E, 1516 m, 8 August 2010; colls: R. W. Sites

& A. Mbogho, shallow silty stream w/ algae & Typha angustifolia, L-1225;, Kindai

Reservoir at Singida, 04º50.065'S 34º44.173'E, 1484 m, 8 August 2010, colls R. W. Sites

& A. Mbogho, tall grasses, L-1226; Singidan Reservoir at Singida, 04º47.568'S

34º44.570'E, 1483 m, 8 August 2010, colls R. W. Sites & A. Mbogho, Typha angustifolia roots & submergent vegetation, L-1227; Mtinko stream at Mtinko, 04º33.525'S

34º50.649'E, 1552 m, 9 August 2010, colls R. W. Sites & A. Mbogho, pooled temp stream w/ algae, L-1229; Mwanga River at Mwanga, 04º17.496'S 34º51.969'E, 1531 m,

9 August 2010, colls R. W. Sites & A. Mbogho, stream pools w/ algae and marginal vegetation, L-1230. TANGA Region: Mazinde stream at Mazinde, 04º48.209'S

38º12.615'E, 433 m, 13 August 2010, colls R. W. Sites & A. Mbogho, narrow stream with tall emergent macrophytes, L-1252; Kwamaluli River at Mbagai, 04º58.561'S

38º23.154'E, 465 m, 14 August 2010, colls R. W. Sites & A. Mbogho, shaded narrow stream w/overhanging veg and undercuts, L-1256.

Laccocoris spurcus congoensis Poisson

(Fig. 9)

Naucoris spurcus Stål, 1856: Ofv. Kong. Vet. Ak. Forh., 13: 197–198.

Laccocoris spurcus: Stål, 1865, Holmiae, p. 178.

Laccocoris spurcus congoensis Poisson, 1949: Inst. Parcs Nat. Congo, 58: 67-69.

Discussion

Laccocoris spurcus is one of the largest species of Laccocoris in Africa. The nominate subspecies is known from South Africa. Poisson (1949) described a subspecies from

Democratic Republic of Congo which is known to have wing polymorphism, with which are associated differences in pronotum shape. Poisson (1949) reported that approximately 90% of the individuals from Democratic Republic of Congo were

26 subbrachypterous or brachypterous and 10% macropterous. All specimens collected from Tanzania were brachypterous.

Diagnosis

Laccocoris spurcus congoensis is distinguished easily from known congeners in Tanzania by its large size (length > 10 mm). Brachypterous individuals have the posterolateral corners of the pronotum narrowly rounded and slightly angled caudad, whereas macropterous individuals have the posterolateral corners of the pronotum broadly rounded, similar to those of L. limigenus. The two subspecies of L. spurcus can be distinguished from each other by the shape of the labrum; it is broadly triangular in L. spurcus congoensis and acutely triangular in the nominate subspecies (Linnavuori 1987), and the posterior spacing of the eyes is slightly less in L. spurcus congoensis (Poisson

1949).

Distribution

L. spurcus congoensis is known from Democratic Republic of Congo, whereas the nominate subspecies was known only from Cape Province in South Africa. In Tanzania,

L. s. congoensis was found in only one locality at high elevation at Mbeya Peak in Mbeya

Region.

Material Examined

TANZANIA: MBEYA Region: Maboo stream at Mbeya Peak village, 08º50.432'S

33º25.060'E, 2433 m, August 2010, colls R. W. Sites & A. Mbogho, vegetated pool of small high elevation stream, L-1207.

Subfamily Naucorinae

Macrocoris flavicollis Signoret

(Fig. 17)

Macrocoris flavicollis Signoret 1860: Soc. Ent. France An., 8: 970.

Discussion

In his treatment on the Hemiptera fauna of Madagascar, Signoret (1860) established the genus Macrocoris with M. flavicollis as the type species. Poisson (1949) described two subspecies of M. flavicollis based on pronotum characteristics and body size. More specifically, M. f. chariensis Poisson is known from only the Chari Basin of Chad and has the pronotum clear yellow with few or no small black spots and is smaller in body size.

M. f. uelei Poisson has the pronotum with larger, confluent black spots and the anterior black spots on the posterior lobe are larger than those of the nominate subspecies. This subspecies was described from Uele in the Democratic Republic of Congo (Poisson

1949) and also is known from Chad (Bebedja), Ivory Coast, central Sudan, and was found among Ceratophyllum and other vegetation and in a swimming pool (Linnavuori

1981). M. f. flavicollis is known from eastern Sudan south throughout most of Africa south of the ranges of M. f. chariensis and M. f. uelei and was recorded from marginal emergent vegetation in pans and a marsh as deep as 2.5 ft in Zimbabwe (Weir 1966).

Linnavuori (1981) reported that this subspecies occurs among vegetation in eutrophic waters.

Of all naucorid species collected in Tanzania, M. flavicollis was the most frequently encountered; collected at 34 of the 127 sites sampled during the entire study

28 period. The species was found in large and small lentic water bodies ranging from lakes to roadside pools and with grasses, algae, lily pads, and usually heavily vegetated. This species was also collected from vegetated margins of sandy and muddy, slow streams.

Water bodies with disturbances including grazing, irrigation or other human activities tended to harbor fewer Macrocoris flavicollis.

Diagnosis

This species is variable in appearance, but fresh specimens have a distinctly green head, pronotum, connexiva and legs, which fade to yellow in alcohol or exposure to light. The pronotum has scattered large, black spots and sometimes small black punctures. At the posterior margin of the head, M. flavicollis has a pair of median black spots which sometimes are contiguous, comprising two vertices more or less elongated anteriorly.

However, the key by Poisson (1959) requires that M. flavicollis have the spots absent or only faintly indicated. The connexiva are concolorous. Body length is 8.9–11.2 mm.

Distribution

In addition to Madagascar and Zanzibar, from which M. flavicollis was described

(Signoret 1860), this species has been recorded from Sudan (Linnavuori 1971, 1981) west through Chad (Poisson 1949, Linnavuori 1981), Central African Republic, Nigeria

(Linnavuori 1981), Benin (Poisson 1951), and Ivory Coast (Linnavuori 1981), south through Uganda, Kenya (Montandon 1914, Poisson 1967), Congo (Poisson 1967,

Linnavuori 1971), Tanzania (Poisson 1963), Rwanda (Poisson 1949), Democratic

Republic of Congo (Poisson 1949, 1963), and Mozambique (Montandon 1899).

The species is widely distributed in Tanzania and was collected in almost every geological region and from greatly varied elevations (30 to 1795 m). Specifically, it was collected from Kagera, Kigoma, Mara, Mbeya, Morogoro, Mwanza, Rukwa, Ruvuma, and Tanga regions. Macrocoris flavicollis was not collected from the central part of

Tanzania (Dodoma and Singida regions) and the neighboring regions of Manyara, Arusha and Kilimanjaro.

Material Examined

TANZANIA: MWANZA Region: Kisesa, ca. 20 km NE Mwanza, 02º33.345'S

33º03.652'E, 1230 m, 24 July 2010, colls R. W. Sites & A. Mbogho, vegetated pond margins, L-1144; near Kashishi, 1344 m, 02º16.239'S 033º48.314'E 24, July 2010, colls

R. W. Sites & A. Mbogho, vegetated pond margins L-1149; Nyanguge, 02º32.619'S

33º13.181'E, 1159 m, 24 July 2010, colls R. W. Sites & A. Mbogho, large vegetated pond, no cattle usage, L-1151. MARA Region: Tairo, ca. 5 km E of Bunda, 02º33.827'S

33º45.695'E, 1230 m, 24 July 2010, colls R. W. Sites & A. Mbogho, heavily vegetated pond margins, L-1146; Ushashi, ca. 5 km N of Nyiendo, 02º00.540'S 33º55.241'E, 1344 m, 24 July 2010, colls R. W. Sites & A. Mbogho, vegetated pond margins, L-1147.

KAGERA Region: ca. 5 km E of Bhiaramulo, 02º38.159'S 31º20.789'E, 1427 m, 26 July

2010, colls R. W. Sites & A. Mbogho, roadside pool with grasses, algae, lily pads, L-

1154; Benako Reservoir, 02º28.081'S 30º51.355'E, 1492 m, 26 July 2010, colls R. W.

Sites & A. Mbogho, marginal vegetation on dam, L-1156; Karenge, 03º07.936'S

31º04.530'E, 1234 m, 27 July 2010, colls R. W. Sites & A. Mbogho, shallow pools with heavy vegetation and cattle use, L-1157. KIGOMA Region: W of Kibondo, Mpemvyi

River, 03º44.028'S 30º41.945'E, 1225 m, 27 July 2010, colls R. W. Sites & A.

Mbogho, pooled areas of slow stream, L-1160; Nyakivaya Lake, ca. 5 km N of Kasulu,

04º16.455'S 30º26.111'E, 1165 m, 27 July 2010, colls R. W. Sites & A. Mbogho, shallow small lake with heavy vegetation, L-1161; Luchugi River, 04º31.733'S

30º09.699'E, 1217 m, 27 July 2010, colls R. W. Sites & A. Mbogho, small stream, L-

1162; 15 km N of Kigoma, Mungonya River, 04º50.337'S 29º40.001'E, 806 m, 28 July

2010, colls R. W. Sites & A. Mbogho, muddy bank with sparse marginal vegetation, L-

1169; Msimba, ca. 18 km N of Kigoma, 04º51.058'S 29º42.552'E, 807 m, 28 July 2010, colls R. W. Sites & A. Mbogho, very slowly flowing stream with marginal vegetation, L-

1170; Katosho Reservoir, 8 km E of Kigoma, 04º51.221'S 29º39.436'E, 802 m, 28 July

2010, colls R. W. Sites & A. Mbogho, emergent dead tall grasses in turbid reservoir, L-

1171; Gombe National Park, Mwagongo, Nkonya River, 04º37.351'S 29º38.319'E, 775 m, 29 July 2010, colls R. W. Sites & A. Mbogho, gravel/rocky clear stream with sparse marginal veg, L-1173; Gombe National Park, Rutanga stream, 04º39.111'S 29º37.660'E,

776 m, 29 July 2010, colls R. W. Sites & A. Mbogho, sandy & muddy stream w/ heavy vegetation, L-1175. RUKWA Region: 93 km N of Mpanda, Kapipula stream,

05º42.233'S 30º41.563'E, 1326 m, 30 July 2010, colls R. W. Sites & A. Mbogho, stream wetland w/ dense emergent grasses, L-1186; 57 km N of Mpanda, Masito /Ogala,

05º56.742'S 30º57.812'E, 1144 m, 30 July 2010, colls R. W. Sites & A. Mbogho, slow stream, coarse roots at margin, L-1187; Sau River at Kasunga, 08º03.444'S 31º30.112'E,

1795 m, 2 August 2010, colls R. W. Sites & A. Mbogho, marginal dense vegetation in cattle pool of stream, L-1200; Kamawe River at Puwi, 08º21.029'S 31º50.054'E, 1598 m,

3 August 2010, colls R. W. Sites & A. Mbogho, clear stream, marginal vegetation, some 31 rocks, L-1205. MBEYA Region: 1 km N of Nyenjele, Muwawa River, 08º55.289'S

32º21.583'E, 1448 m, 3 August 2010, colls R. W. Sites & A. Mbogho, pool with overhanging grasses, L-1206; Itunga inlet, Lake Nyassa, 09º35.094'S 033º56.769'E, 482 m, 4 August 2010, coll R. W. Sites and A. Mbogho, heavy marginal vegetation at boat ramp, L-1208. RUVUMA Region: 20 km N of Songea, Lumecha River at Samangula,

10º30.149'S 35º40.268'E, 994 m, 5 August 2010, colls R. W. Sites & A. Mbogho, overhanging grasses, L-1211; Hanga River at Ngadinda, 10º10.264'S 35º39.611'E, 777 m, 5 August 2010, colls R. W. Sites & A. Mbogho, overhanging veg and matete, L-1213.

MOROGORO Region: Mikumi National Park, 07º21.608'S 37º03.337'E, 529 m, 7

August 2010, colls R. W. Sites & A. Mbogho, heavily vegetated seasonal pond, L-1219; ca. 21 miles NW of Morogoro, 06º29.634'S 37º33.465'E, 372 m, 7 August 2010, colls R.

W. Sites & A. Mbogho, heavily vegetated pond ringed with reed grass, L-1220.

SINGIDA Region: Kipumbuiko River, 05º05.667'S 34º45.940'E, 1516 m, 8 August 2010; colls: R. W. Sites & A. Mbogho, shallow silty stream w/ algae & Typha angustifolia, L-

1225; TANGA Region: river from Soni to Mombo, 2 km E of Vuga Junction,

04º52.040'S 38º20.906'E, 460 m, 14 August 2010, colls R. W. Sites & A. Mbogho, grassy overhangs in rocky streams, L-1253; Kwamaluli River at Mbagai, 04º58.561'S

38º23.154'E, 465 m, 14 August 2010, colls R. W. Sites & A. Mbogho, shaded narrow stream w/ overhanging vegetations and undercuts, L-1256. ZANZIBAR: KASKAZINI

Region: Kitope River at Zingwezingwe Bridge, 06º00.963'S 39º14.734'E, 39 m, 17

August 2010, R. W. Sites & A. Mbogho, discontinuous pools w/ aquatic veg and grasses,

L-1265; Kitope River at Mkaratini Bridge, 06º00.714'S 39º14.841'E, 30 m, 17 August

2010, R. W. Sites & A. Mbogho, discontinuous pools w/ aquatic veg and grasses, L-1266 32

Macrocoris sp.

(Fig. 18)

Discussion

This genus is endemic to the African continent and is represented by 10 species, 3 of which are endemic specifically to Madagascar. Macrocoris and its sister genus

Neomacrocoris, also endemic to Africa, are relatively dorsoventrally robust for the family. Both genera occupy heavily vegetated margins of water bodies. As a generality,

Macrocoris tends to be found more in lentic water bodies, whereas Neomacrocoris tends to occur in marginal vegetation of slow streams. Macrocoris can be distinguished from

Neomacrocoris by its wider head, negligible claval commissure, and well-developed parameres of the male genitalia.

This appears to be an undescribed species of Macrocoris. Species in this genus are distinguished based on characteristics of pronotum punctation and maculation, connexiva coloration, and characters related to male genitalia and medial lobes of the 8th abdominal tergum. Because this species does not agree in these features with any described species or subspecies, I consider it to be undescribed. The most recent treatment of this genus

(Poisson 1959) did not provide a rigorous assessment of interspecific distinctions.

Although Poisson described many species and subspecies of both Macrocoris and

Neomacrocoris, his work has been found to be unreliable. Thus, generic revision is necessary and description of this new species should be included therein.

Diagnosis

The pronotum of this species is beset with dense dark punctation and the body is highly convex dorsally. Body length is 10.0–10.9. Poisson (1959) presented a key to the species and subspecies of Macrocoris, in which he separated the taxa into two groups based on connexival color patterns. One group is concolorous and the other has the posterolateral corners black or densely maculated. As such, this undetermined species falls into the latter group, which includes M. convexus, M. nigropunctatus Montandon, M. distinctus Bergroth, and M. transvaalensis Distant.

Distribution

This species was found in only one locality and was among marginal vegetation on a dam in Kagera Region.

Material Examined

TANZANIA: KAGERA Region: Benako Reservoir, 02º28.081'S 30º51.355'E, 1492 m,

26 July 2010, colls R. W. Sites & A. Mbogho, marginal vegetation on dam, L-1156.

Naucoris obscuratus Montandon

(Fig. 6)

Naucoris obscuratus Montandon 1913b: Bull. Sect. Sci. Acad. Roum. 1: 221.

Discussion

This species is common in eutrophic rivers and normally is found among vegetation and debris (Linnavuori, 1971) in rivers, lakes, and pools (Linnavuori, 1981). Naucoris obscuratus is the only member of the genus Naucoris known from Tanzania. Even

34 though N. obscuratus has been recorded from throughout Africa, in this study it was collected at only a few localities in Tanzania and most were from the northern part of the country. Montandon (1913b) listed this species from six localities, including “Hego

Tanganika”, which Poisson (1967) repeated the Montandon localities and referred to the

Tanganyika record as “lac Tanganyika”. Because Lake Tanganyika borders four countries, this record was not specific to a particular country. Thus, the collections reported here represent the first record of N. obscuratus from Tanzania.

Diagnosis

Naucoris obscuratus can be distinguished from the other two genera of Naucorinae in

Tanzania by its shape and size. N. obscuratus is much flatter than the dorsally convex

Macrocoris and Neomacrocoris and body length is only 6.5 to 8.7 mm. Further, males do not have tergites 7 and 8 modified, whereas those of the other genera have medial lobes with modifications that are species specific. Males have parameres that are well developed and overlapping, whereas Macrocoris has parameres that do not overlap and

Neomacrocoris lacks parameres.

Distribution

In the original description, Montandon (1913b) reported the distribution as from Sudan to

Lake Tanganyika, Egypt, Nigeria, and South Africa, and this was largely repeated by

Poisson (1967). The East African countries of Uganda and Kenya were added by

Montandon (1914) and Poisson (1963), respectively. Congo was added in the description of the subspecies N. o. nanus (Poisson 1960a). West Africa was added (Linnavuori,

1971) and was followed by specific country records for Ivory Coast, Ghana, Nigeria, and

Chad (Linnavuori, 1981). During this study, N. obscuratus was found mostly in northern

Tanzania. More specifically, it was collected from seven localities in Kagera, Kigoma,

Kilimanjaro, Mara and Pwani regions.

Material Examined

TANZANIA: KAGERA Region: Benako Reservoir, 02º28.081'S 30º51.355'E, 1492 m,

26 July 2010, colls R. W. Sites & A. Mbogho, marginal vegetation on dam, L-1156.

KIGOMA Region: Nyakivaya Lake, ca. 5 km N of Kasulu town, 04º16.455'S

30º26.111'E, 1165 m, 27 July 2010, colls R. W. Sites & A. Mbogho, shallow small lake with heavy vegetation, L-1161. KILIMANJARO Region: Jiwemawe Reservoir at

Ndungu, 04º25.496'S 38º05.201'E, 515 m, 13 August 2010, colls R. W. Sites & A.

Mbogho, vegetated margins of reservoir & below spillway, L-1251. MARA Region:

Tairo, ca. 5 km E of Bunda, 02º33.827'S 33º45.695'E, 1230 m; 24 July 2010, colls R. W.

Sites & A. Mbogho, heavily vegetated pond margins, L-1146; Ushashi, ca. 5 km N of

Nyiendo, 02º00.540'S 33º55.241'E, 1344 m, 24 July 2010, colls R. W. Sites & A.

Mbogho, vegetated pond margins, L-1147. PWANI Region: pond by Ruvu River W of

Bagamoyo, 06º28.545'S 38º50.614'E, 2 m, 16 August 2010, colls R. W. Sites & A.

Mbogho, water lettuce and other vegetation, L-1263.

Neomacrocoris bondelaufa Sites

(Figs. 10, 21)

Neomacrocoris bondelaufa Sites & Mbogho 2012: Zootaxa (in press).

Discussion

Neomacrocoris bondelaufa is a newly described species and was first found during these collections. Although the species is distributed widely throughout the Rift Valley of western Tanzania, its distinction from its congeners of the region apparently was overlooked because of its superficial similarity. During this project, N. bondelaufa was collected syntopically with N. handlirschi at L-1202 and with N. parviceps at L-1157,

1164, 1182, and 1202. All three species were collected together in the Msanzi River in

Rukwa Region (L-1202). The habitat in which these species occur is the densely vegetated margin of streams where current is minimal. Typically, specimens were collected in undercuts and dense vegetative cover with muddy substrate.

Diagnosis

N. bondelaufa is nearly indistinguishable superficially from several congeners, with which it co-occurs, including N. parviceps and N. handlirschi, without examination of male genitalia. However, fresh specimens of N. parviceps are reddish-orange on the abdominal sterna, whereas N. bondelaufa and N. handlirschi are brown. N. handlirschi generally is only slightly larger than N. bondelaufa, but the size ranges overlap. Until other diagnostic characters are discovered, females must be associated with males for identification using male genitalia. Specifically, the apex of the phallosoma has a strong gibbosity to the right and ventrally in N. bondelaufa and directly to the right in N. handlirschi, whereas it has an elongate flange to the right in N. parviceps. Further, the left side of the shaft of the phallosoma beyond the gibbosity is arcuate and the left side of the apex appears flattened in dorsal view in N. bondelaufa, whereas the shaft is straight

37 and the flattening is on the distal margin in N. handlirschi. The apical flattening appears flattened on the left side in dorsal view in N. parviceps. Body length is 8.2–9.2 mm.

Distribution

This species is distributed widely throughout the Rift Valley of western Tanzania and also occurs in South Africa.

Type Material Examined

HOLOTYPE ♂: TANZANIA: RUKWA Region: Sau River at Kasunga, 08º03.444'S

31º30.112'E, 1795 m, 2 August 2010, colls R. W. Sites & A. Mbogho, marginal dense vegetation in cattle pool of stream, L-1200. Paratypes: same data as holotype (18♂,

21♀). Rukwa Region, Kapipula stream, 93 km N of Mpanda, and 05 º 42.233’S

30º41.563'E, 1326 m, 30 July 2010, colls R. W. Sites & A. Mbogho, stream wetland w/ dense emergent grasses, L-1186 (5♂, 8♀). KIGOMA Region: Kazuramimba,

05º00.155'S 30º00.971'E, 1082 m, 30 July 2010, colls R. W. Sites & A. Mbogho, muddy pools of small stream with marginal vegetation, L-1182 (4♂).

Additional Material Examined

TANZANIA: KAGERA Region: Karenge, 03º07.936'S 31º04.530'E, 1234 m, 27 July

2010, colls R. W. Sites & A. Mbogho, shallow pools with heavy vegetation and cattle use, L-1157. KIGOMA Region: W of Kibondo, Mpemvyi River, 03º44.028'S

30º41.945'E, 1225 m, 27 July 2010, colls R. W. Sites & A. Mbogho, pooled areas of slow stream, L-1160; Luchugi River, 04º31.733'S 30º09.699'E, 1217 m, 27 July 2010, colls R.

W. Sites & A. Mbogho, small stream, L-1162; tributary to Kalembela River, 04º40.743'S

30º03.994'E, 1244 m, 27 July 2010, colls R. W. Sites & A. Mbogho, irrigation canal, L- 38

1163; Kalembela River, 04º40.813'S 30º03.964'E, 1249 m, 27 July 2010, colls R. W.

Sites & A. Mbogho, marginal vegetation in slow current, L-1164. MARA Region:

Ushashi, ca. 5 km N of Nyiendo, 02º00.540'S 33º55.241'E, 1344 m, 24 July 2010, colls

R. W. Sites & A. Mbogho, vegetated pond margins, L-1147. MBEYA Region: 1 km N of Nyenjele, Muwawa River, 08º55.289'S 32º21.583'E, 1448 m, 3 August 2010, colls R.

W. Sites & A. Mbogho, pool with overhanging grasses, L-1206. RUKWA Region: 110 km N of Mpanda, 05º37.745'S 30º36.039'E, 1295 m, 30 July 2010, colls R. W. Sites & A.

Mbogho, shallow creek w/ muddy bottom and marginal grasses, L-1185; 57 km N of

Mpanda, stream btw Masito & Ogala, 05º56.742'S 30º57.812'E, 1144 m, 30 July 2010, colls R. W. Sites & A. Mbogho, slow stream, coarse roots at margin, L-1187; 20 km SW of Sumbawanga, Katuka River at Katuka, 08º06.641'S 31º31.786'E, 1858 m; 2 August

2010, colls R. W. Sites & A. Mbogho, pools of stream and narrow channel w/ dense vegetation; L-1201; Msanzi River at Msanzi, 08º11.411'S 31º31.295'E, 1778 m; 2 August

2010, colls R. W. Sites & A. Mbogho, rocks, mud, gravel, sand, marginal grasses, L-

1202; Kamawe River at Puwi, 08º21.029'S 31º50.054'E, 1598 m; 3 August 2010, colls R.

W. Sites & A. Mbogho, clear stream, marginal vegetation, some rocks, L-1205. SOUTH

AFRICA: Swaziland, Malolotja, Res. Dam at Camp, 7-4-97 / 26o 09’S 31o 08’E, 1800 m.a.s.l. (1♂).

Neomacrocoris handlirschi (Montandon)

(Figs. 11, 25)

Macrocoris handlirschi Montandon 1909: Bul. Soc. Rom. Stiinte 18: 55.

Neomacrocoris handlirschi: Montandon 1913a, Bul. Soc. Rom. Stiinte 22: 332.

Discussion

This species originally was described as Macrocoris handlirschi from southern South

Africa (Montandon 1909). Four years later, this and two other species of Macrocoris were transferred to the new genus Neomacrocoris by Montandon (1913a). Montandon

(1909) described the species from a series but did not designate a type specimen. Sites and Mbogho (2012) located a single syntype in the Naturhistoriches Museum in Vienna and designated it as the lectotype. Poisson (1948) reported that the species has a wide size range of 8.5–10 mm length and 5.2–5.3 mm width. N. handlirschi was distinguished from N. angusticeps by Montandon (1914) by its smaller size; and the pronotum with more distinct black punctation, posterolateral corners blunter and narrowly rounded, and a superficial transverse furrow of the pronotal disc. Poisson (1948) also noted a slight indication of a posterior transverse furrow of the pronotal disc.

The variety N. handlirschi var. armatus was described by Poisson (1948) from seven localities in the Belgian Congo (Luebo, Matadi, Boma, Luluabourg, Mayidi, Haut

Uelé, and Katanga) and distinguished from the nominate form by the condition of the 7th abdominal tergum. Specifically, the left lobe had 1–8 denticles in n. handlirschi armatus, whereas it was without denticles in the nominate subspecies. In a population sampled from a small stream in Gombe National Park in northwestern Tanzania were males with from zero to five denticles on the left medial lobe of the 7th abdominal tergum, thereby presenting characteristics of both subspecies. Because both character states were present in the same population, the subspecific designation based solely on presence or absence of denticles is not valid; thus, Sites and Mbogho (2012) synonymized the subspecies.

Diagnosis

This species may be distinguished from all of its congeners except N. ndugai, known from only Rwanda, by the slight flattening of the pronotum and by the presence of short, thick setae sparsely distributed over the surface of the scutellum and hemelytra. In fresh specimens, the short setae can be found sparsely over the entire dorsal surface of the hemelytra, including the clavus and membrane; however, the hairs apparently are fragile and less evident on older or mishandled specimens. The condition of the apex of the phallosoma distinguishes N. handlirschi from N. ndugai. The female subgenital plate is gently sinuate to the distal spatulate process, which differs from only N. vuga among the known congeneric females from Tanzania. Body length is 8.6–9.2 mm.

Distribution

The species occurs from eastern Sudan (Linnavuori 1981) and the Democratic Republic of Congo and Uganda south to South Africa. As such, N. handlirschi and N. parviceps might have the widest distribution of any species of Neomacrocoris. In Tanzania, N. handlirschi was reported from the Usambara Mountains in the northeastern part of the country (Poisson 1963). N. handlirschi was collected syntopically with N. bondelaufa at

L-1202 and with N. parviceps at L-1174 and 1202. All three species were collected together in the Msanzi River in Rukwa Region (L-1202). Specimens were collected in densely vegetated stream margins with muddy or gravelly substrate.

Material Examined

TANZANIA: KIGOMA Region: Gombe National Park, Mitumba stream, S 04º 38.405'

E 029º 37.823', 776 m, 29 July 2010, colls R. W. Sites & A. Mbogho, heavily-vegetated

41 narrow forest stream, L-1174. RUKWA Region: Msanzi River at Msanzi, S 08º 11.411',

E 031º 31.295', 1778 m, 2 August 2010, colls R. W. Sites & A. Mbogho, rocks, mud, gravel, sand, marginal grasses, L-1202.

Neomacrocoris karimii Poisson

(Figs. 12, 23)

Neomacrocoris karimii Poisson 1963: Bull. Inst. Franc. Afr. Noire 25A: 1173, 1189-

1191.

Discussion

This species is known from only a single male specimen collected in a small stream over rocks near the route of Muheza to Amani on the road to the Karimi Tea

Estate in the Usambara Mountains of Tanzania, as was given in the original description.

No other account of this species has been published besides its listing in the La Rivers

(1971) catalog and recent inclusion in the revision of the genus (Sites and Mbogho,

2012). During this study, this species was not collected despite collecting at the type locality and in nearby streams. The substrate of the stream running through the tea plantation is covered heavily with red silt resulting from the agricultural activities, and very few insects were found overall. Elsewhere in the Usambara Mountains, a superficially similar species (see N. vuga below) was collected, which I was able to determine as distinct only after examination of the genitalia of N. karimii (Fig. 23). The type specimen of N. karimii is card-mounted and housed in the Muséum National d'Histoire Naturelle of Paris, but is missing the posterior abdominal segments. Two microscope slides containing the aedeagus and 7th and 8th segments of this specimen are

42 housed in the Drake Collection in the U. S. National Museum (USNM). Poisson (1963) illustrated the body, aedeagus, and 7th and 8th terga.

Diagnosis

This species is distinguished from its congeners in Tanzania by its ovate body (Fig. 12) and the distinct male phallosoma (Fig. 23). Because this species is known from only the holotype, the structure of the female subgenital plate is not known. Body length is 10.3 mm.

Distribution

This species is known from only a single male specimen collected from Tanga Region, near the route of Muheza to Amani on the road to the Karimi Tea Estate in the Usambara

Mountains of Tanzania.

Material Examined

HOLOTYPE: TANGANYIKA, ruisselet de karini /type /Neomacrocoris karinii Poiss.

(NMHN, ♂ body; USNM, slide-mounted abdominal segments 7– 8 and phallosoma).

Neomacrocoris parviceps (Montandon)

(Figs. 13, 14, 22, 26)

Macrocoris parviceps Montandon 1900: Terme. Fuz. 23: 422.

Neomacrocoris parviceps Montandon: 1913a: Bul. Soc. Rom. Stiinte, 22: 332.

Discussion

This is arguably the most common species of Neomacrocoris in Africa. Poisson (1948) considered Neomacrocoris parviceps to resemble N. handlirschi and N. nanus, and is

43 variable in pigmentation and size. It has been given as 7.8 (Montandon 1900) to 9 mm

(Poisson 1948) in length. The specimens from throughout Tanzania range from 7.36 to

8.56 mm. This species is variable in expression of pigmentation, size, as well as other subtle differences in degree of angle and spination of abdominal tergal lobes and curvature in the structure of the phallosoma. Poisson (1948) described a variety (N. parviceps var. ocellatus) based on more pronounced punctation near the margins of the pronotum and a more clear (“nette”) claval commissure than that of N. p. parviceps, but gave no distribution information. Although he treated it as a variety, Poisson referred to the alternate condition as N. parviceps parviceps, implicitly conferring subspecific rank to ocellatus. In later papers, he referred to it more formally as a subspecies (N. p. ocellatus) (Poisson 1949) and reported it from Maua, Kilimanjaro Region, Tanganyika

(Poisson 1963). Despite collecting throughout the area of Maua, no naucorid specimens were found.

Diagnosis

This is a moderately sized species with an obviously reddish-orange abdominal sternum in living specimens. Thick setae on the hemelytra are restricted to the embolium and adjacent corium. The longitudinal flange and apex of the phallosoma readily distinguish this species. The female subgenital plate is gently sinuate to the distal spatulate process.

Body length is 7.36–8.56 mm.

Distribution

The type locality is Dar Es Salaam, Tanzania and the lectotype is housed in the

Hungarian Natural History Museum - Budapest. This species is known from Sudan

(Linnavuori 1981) and was considered to be widespread in East Africa (Poisson 1948).

This species also has been recorded from Ghana, Ivory Coast, Upper Volta, Democratic

Republic of Congo, Uganda, and South Africa (Sites and Mbogho, 2012). This species was collected from throughout Tanzania, including the island of Zanzibar.

Material Examined

TANZANIA: KAGERA Region: ca. 5 km E of Bhiaramulo, 02º38.159'S 31º20.789'E,

1427 m, 26 July 2010, R. W. Sites & A. Mbogho, roadside pool with grasses, algae, lily pads, L-1154; Karenge, 03º07.936'S 31º04.530'E, 1234 m, 27 July 2010, R. W. Sites &

A. Mbogho, shallow pools with heavy vegetation and cattle use, L-1157. KIGOMA

Region: Kalembela River, 04º40.813'S 30º03.964'E, 1249 m, 27 July 2010, R. W. Sites

& A. Mbogho, marginal vegetation in slow current, L-1164; Gombe National Park,

Mitumba stream, 04º38.405'S 29º37.823'E, 776 m, 29 July 2010, R. W. Sites & A.

Mbogho, heavily vegetated narrow forest stream, L-1174; Kazuramimba, 05º00.155'S

30º00.971'E, 1082 m, 30 July 2010, R. W. Sites & A. Mbogho, muddy pools of small stream with marginal vegetation, L-1182. MBEYA Region: Tanganyika, Maji Moto,

3,700 ft., 33o 40’ E, 8o 38’ S, 9.viii.1959, Cambridge E. African Exped., B.M. 1960-50

(BMNH, 1♂). RUKWA Region: ca. 54 km E. of Karema, 06º37.275'S 30º39.593'E,

1086 m, 31 July 2010, R. W. Sites & A. Mbogho, shaded stream, negligible flow, L-

1193; Mpanda, Kasimba River, 06º21.602'S 31º04.382'E, 1064 m, 1 August 2010, R. W.

Sites & A. Mbogho, grasses hanging into river, L-1195; Msanzi River at Msanzi,

08º11.411'S 31º31.295'E, 1778 m, 2 August 2010, R. W. Sites & A. Mbogho, rocks, mud, gravel, sand, marginal grasses, L-1202; Tanganyika, Rukwa Valley, 9 xi 1963, E.S.

Brown, 536, B.M. 1966-102 (BMNH, 3♂, 2♀). RUVUMA Region: 20 km N of Songea,

Lumecha River at Samangula, 10º30.149'S 35º40.268'E, 994 m, 5 August 2010, R. W.

Sites & A. Mbogho, overhanging grasses, L-1211. TANGA Region: river from Soni to

Mombo, 2 km E of Vuga Junction, 04º52.040'S 38º20.906'E, 460 m, 14 August 2010, R.

W. Sites & A. Mbogho, grassy overhangs in rocky stream, L-1253; Kwamaluli River at

Mbagai, 04º58.561'S 38º23.154'E, 465 m, 14 August 2010, R. W. Sites & A. Mbogho, shaded narrow stream w/ overhanging veg and undercuts, L-1256; Kihuuwi River at

Timba, 05º07.683'S 38º41.418'E, 174 m, 15 August 2010, R. W. Sites & A. Mbogho, marginal grasses with silty/rocky bottom, L-1260; Jembe stream at Muheza, 05º10.260'S

38º47.106'E, 182 m, 15 August 2010, R. W. Sites & A. Mbogho, grassy overhangs in urban stream, L-1261. ZANZIBAR: KASKAZINI Region: Kitope River at

Zingwezingwe Bridge, 06º00.963'S 39º14.734'E, 39 m, 17 August 2010, R. W. Sites &

A. Mbogho, discontinuous pools w/ aquatic veg and grasses, L-1265; Kitope River at

Mkaratini Bridge, 06º00.714'S 39º14.841'E, 30 m, 17 August 2010, R. W. Sites & A.

Mbogho, discontinuous pools w/ aquatic veg and grasses, L-1266. ZANZIBAR:

KUSINI Region: Mwera River at Mwera, 06º08.811'S 39º16.264'E, 23 m, 17 August

2010, R. W. Sites & A. Mbogho, flooded vegetated field margin, L-1267.

Neomacrocoris usambaricus Montandon

(Figs. 15, 19)

Neomacrocoris usambaricus Montandon 1913a: Bull. Soc. Rom. Stiinte, 22: 332-334.

Neomacrocoris usambaricus: Sites & Mbogho 2012, Zootaxa (in press) (neotype designation).

Discussion

This is the type species of the genus as designated by Štys and Jansson (1988). The type specimen was collected in the western Usambara Mountains of Tanzania. Although the holotype has been lost since the 1930s, a neotype from the Democratic Republic of

Congo was designated by Sites and Mbogho (2012). Despite collecting several streams in that area, nothing resembling the neotype was found.

Diagnosis

Although it is large, N. usambaricus is not in the group of distinctly large and ovate species. The pronotum is largely devoid of dark maculation. Currently, only the female is known and the subgenital plate is gently sinuate to the distal spatulate process. Body length of the neotype is 10.96 mm.

Material Examined

NEOTYPE: [DEMOCRATIC REPUBLIC OF CONGO]: Tuevo, 2-VIII-1911, R.

Mayné / R. det., A, 202 / Neomacrocoris usambaricus 1913 Montandon, det Montandon

1914 (RMCA, ♀).

Neomacrocoris vuga Sites

(Figs. 16, 20, 24)

Neomacrocoris vuga Sites & Mbogho 2012: Zootaxa (in press).

Discussion

This is a newly described species and was first found during these collections. This species occurred syntopically with N. parviceps at the type locality. Sweeping dense submerged vegetation along the margins of a small stream yielded nine adults and many immatures at the type locality and another site further upstream approximately 600 m higher in the mountains. This species superficially is similar to N. karimii, which is known from the same general area of the Usambara Mountains in Tanzania, and N. vaneyeni from the Democratic Republic of Congo. This species is named for Vuga

Junction in the Usambara Mountains near Mombo, adjacent to the stream in which it was collected.

Diagnosis

This species is large and ovate as are N. karimii and N. vaneyeni (from Democratic

Republic of Congo), but can be diagnosed readily by the male genitalia. Specifically, the apex of the phallosoma is strongly and sharply hooked to the left, and a subapical rounded lobe extends dorsally and to the left (Fig. 24). The female subgenital plate is broad and has a strong mid-lateral projection before narrowing to the distal spatulate process (Fig. 20). Body length is 10.8–11.9 mm.

Distribution

This species was found at two locations in the river from Soni to Mombo in Usambara

Mountains. The habitat in which it occurred was grassy overhangs and undercuts in a rocky stream with moderate current.

Type Material Examined

HOLOTYPE ♂: TANZANIA: TANGA Region; river from Soni to Mombo, 2 km E of

Vuga Junction, 04º 52.040'S, 38º 20.906'E, 460 m, 14 August 2010, colls R. W. Sites &

A. Mbogho, grassy overhangs in rocky stream, L-1253. PARATYPES, same data as holotype (1♂, 2♀); river from Soni to Mombo, Karense, 1 km E of Soni, 04º 50.284'S,

38º 22.355'E, 1051 m, 14 August 2010, colls R. W. Sites & A. Mbogho, vegetated overhangs and undercuts, L-1254 (1♂, 4♀).

Additional Material Examined

TANZANIA: Tanga Region, W. Usambara Mts. 38 22E, 4 44S, 10 Aug 1996, 5100 ft,

P. Cresswell, creek (Carnegie Museum, 1♀).

Distribution Overview

Naucorids were collected at 78 of the 127 collections (61.4%). Of the 14 species of

Naucoridae now known from Tanzania, Macrocoris flavicollis was the most widely distributed as it was collected in the greatest number of samples (34) and that it was taken in all six land resource zones (Table 3, Fig. 1). The Highlands land resource zone harbored the greatest number of species (12); all but Ctenipocoris africanus and

Neomacrocoris parviceps have been recorded from there. From the collections reported here, two species each were found in only one locality (Laccocoris spurcus congoensis and Macrocoris sp.), and two others not collected in this project each have been recorded from only one locality (Neomacrocoris karimii and N. usambaricus). Six species were recorded only from the East African Rift Valley, suggesting a level of endemism and degree of restriction in the regional fauna. These species include Neomacrocoris

49 bondelaufa (so named for the Rift Valley), N. handlirschi, Macrocoris sp., Aneurocoris insolitus, and Laccocoris limicola.

Table 3. Taxa of Naucoridae found in Tanzanian Land Resource Zones. Number of samples from each zone appears in parentheses below the name of the zone.

* Species that were not collected in this study but are known from Tanzania.

Discussion

Africa harbors a diverse and relatively unstudied fauna in comparison to those of other continents. The faunas of Australia, Europe, and North America have been studied heavily over many years and although discovery of new taxa is ongoing, the proportion

50 of known to unknown species is thought to be much higher than it is for Africa. Fewer and fewer new species are being discovered in temperate regions. Conversely, relatively little is known of the fauna of tropical regions, including Africa; this disproportionate level of knowledge is due to several factors. Much of Africa is tropical, although the northern and southern limits of the continent are temperate. The number of species increases with proximity to the equator, and tropical forests are thought to hold more than

50% of all species (Lovejoy 1997). As such, the Heteroptera fauna of Africa can be generalized to be substantially greater than that of similar-sized temperate land masses.

For the Naucoridae alone, the primarily temperate landmasses or regions have low numbers of species: Australia, 5; Nearctic, 29; Palearctic, 6; whereas the number of species in primarily tropical land masses is staggeringly higher: Neotropical 186;

Oriental, 78; Afrotropical, 71. Despite the higher number of known species for Africa, little faunistic research on the Naucoridae has been conducted, and not since the middle

1900s. This research revealed double the number of species occurring in Tanzania than was previously known. Tanzania is a relatively arid country in tropical Africa, yet the naucorid fauna was found to be greater than that of Australia, Europe, and all of

Palearctic Asia combined. Further these new species were discovered on the east and west sides of the country. Given this high diversity for a mostly arid African country and that three new species were discovered, the diversity and number of undescribed species of Naucoridae in wetter tropical African countries is predicted to be dramatically higher.

Most areas of Africa have not been sampled for naucorids at all, let alone sufficiently.

Areas that have received moderate collecting are South Africa, Sudan, several national parks such as Parc National Albert in Uganda, and now also Tanzania. Many species

51 were described from what is now the Democratic Republic of Congo, a large area of tropical forest in the heart of Africa, but most species are known by only one or a few specimens. Unfortunately, DRC is not conducive for foreign scientists because of political instability, civil chaos, and the associated inherent danger. When the situation becomes more amenable for scientists, this is a region of Africa that should be high on the list for faunistic exploration because it holds great promise of harboring many undescribed taxa. Specifically in Tanzania, potential regions that should be targeted for undiscovered species include the Uldizungwa Mountains, Usambara Mountains,

Ngorongoro Crater, and Mt. Kilimanjaro.

Political instability is among the obstacles for foreign scientific exploration in much of

Africa. Governments of countries that are stable and interested in learning their fauna should reach out to qualified systematic authorities on taxa of interest by providing unfettered access to collecting and export permits. Enabling these researchers to conduct field work that will result in the determination of the local fauna is the only way in which developing countries will be able to understand the composition of their faunas. In addition, providing scholarships to promising African students to study in “western” universities will provide the education and perspective from which to build a human scientific infrastructure that can work effectively to determine the fauna independently and with visiting scientists.

The study presented here represents a great deal of field work and taxonomic investigation because many of these species were known by only a few specimens in various European, African, and American museums. As such, this study is the first of its kind in providing an authoritative identification resource to determine the entire fauna of

Naucoridae of an African country using modern techniques. Publication of this study can serve as a model for other African countries to emulate to report the fauna of Naucoridae or any other group of insects. It is only with resources such as this that scientists and lay people will be able to acquire insight into and develop an understanding and appreciation of their local natural world.

CHAPTER 4 TAXONOMIC KEY

Two subfamilies of Naucoridae (Laccocorinae and Naucorinae), each with 3 genera, were found during this study. The Tanzanian fauna is now known to include 14 species, one of which is represented by two subspecies. In this chapter, a taxonomic key for identification of the species of Naucoridae in Tanzania is presented. Note that the key includes all species that were collected during this study and two species that were not collected but have been reported in the literature.

Key to the species of Naucoridae of Tanzania

1. Front of head folded posteroventrally (Fig. 2). Foreleg pretarsus of females with two

claws. Males with well developed tomentose patch on mesotibia (on females weakly

developed) ………………………………….…………...... ……... Laccocorinae …..2

1’. Front of head not folded posteroventrally (Fig. 3). Foreleg pretarsus of females with

a single claw. Without tomentose patch on mesotibia………...... ….Naucorinae…. 6

2. Hemelytra without veins, embolar and claval sutures absent (Fig. 4) (Lake

Tanganyika) ………………………..……...... ….Aneurocoris insolitus Montandon

2’. Hemelytra with veins, embolar and claval sutures distinct (Fig. 5)……...... ……..3

3. Pronotum with greatest width approximately twice its length at midline (Fig. 5).

Females with two protarsal segments …...... ……....Ctenipocoris africanus Poisson

3’. Pronotum with greatest width approximately three times its length at midline (Fig.

7–9). Females with one protarsal segment ……….…...... Laccocoris.....… 4

4. Pronotum with posterolateral corners broadly rounded or truncated, not produced

caudad (Fig. 8). Hemelytral membrane broad……...... Laccocoris limigenus Stål

4’. Pronotum with posterolateral corners at least slightly produced caudad (Figs. 7, 9).

Hemelytral membrane narrow……...... ……....………….……………………..5

5. Body large, length 10–13 mm……...... Laccocoris spurcus congoensis Poisson

5’. Body small, length <10 mm………………...... …..….. Laccocoris limicola Stål

6. Body elongate and flattened (Fig. 6); male with tergites 7 & 8 unmodified;

parameres well developed; aedeagus without sclerotized vesica distally……………..

……….……...... ………………………………..Naucoris obscuratus Montandon

6’. Body ovate and dorsoventrally robust (Figs. 10–18); male with tergites 7 & 8

modified with asymmetrical medial lobes; parameres present or absent; aedeagus

modified with sclerotized vesica distally...…………………...……………….…... 7

7. Greatest head width across the eyes less than half the greatest pronotum width (Figs.

10–16). Male parameres absent ……………...... ……...... …Neomacrocoris ….8

7’. Greatest head width across the eyes more than half the greatest pronotum width

(Figs. 17–18). Male parameres well developed . …….…...... …....Macrocoris..…15

8. Body large and ovate; length ≥ 10.0 mm; width/length ratio usually ≥ 0.675……… 9

8’. Body smaller and more elongate; length < 10.0 mm; width/length ratio <0.675...…12

9. Females (female of N. karimii unknown) ………………….……...... …….… 10

9’. Males (males of N. usambaricus unknown) …….………………...... ……….… 11

10. Subgenital plate with lateral margins smoothly sinuate to distal spatulate process

(Fig. 19) ...... ……………...... …….. Neomacrocoris usambaricus Montandon

10’. Subgenital plate with strong mid-lateral projections and distal spatulate process (Fig.

20)..……………………………………..……...... ……….Neomacrocoris vuga Sites

11. Phallosoma with strong acute apical hook projecting to left (Fig. 24).…………….

…………………………………………………………. Neomacrocoris vuga Sites

11’. Phallosoma with left side of apex rounded (Fig. 23).Neomacrocoris karimii Poisson

12. Thick setae scattered sparsely on scutellum and hemelytra, including on the clavus,

embolium, corium, and membrane (Fig. 11). Pronotum slightly flattened (Fig.

25)………………………………………..Neomacrocoris handlirschi (Montandon)

12’. Thick setae restricted to embolium and adjacent areas of corium, although thin hair-

like setae can be present throughout (Figs. 10, 13, 14). Pronotum dorsoventrally

robust (Fig. 26)…………………………………………………………………….. 13

13. Lateral margin of embolium slightly less arcuate in anterior 1/3 to ½ (Fig. 10).

Phallosoma with distinct gibbosity on right side (Fig. 21) ......

...... ……………………………………..……...... Neomacrocoris bondelaufa Sites

13’. Lateral margin of embolium evenly arcuate throughout (Figs. 13, 14). Phallosoma

with elongate flange on right side (Fig. 22) …………………………………...… 14

14. Pronotum with black spots on lateral 1/3 larger than spots on central 2/3 (Fig. 13)....

...... ………………Neomacrocoris parviceps ocellatus Poisson

14’. Pronotum with black spots on lateral 1/3 subequal to spots on central 2/3 (Fig. 14)…

…….………..…...……...... Neomacrocoris parviceps parviceps (Montandon)

15. Pronotum mostly yellow (green in live specimens) with sparse large black spots,

some specimens with small dark punctation in addition to larger dark spots (Fig. 17).

Lateral margins of connexiva continuously greenish-yellow…......

………………………………………….…………...Macrocoris flavicollis Signoret

15’. Pronotum with profuse small black spots (Fig. 18). Connexiva with posterolateral

corners dark brown, yellow in anterior ¾………...... ……………… Macrocoris sp.

Figures 2–3. Head showing the difference between subfamilies. (2) Laccocorinae (Laccocoris limigenus) with front folded posteroventrally, (3) Naucorinae (Neomacrocoris parviceps) not folded.

Figures 4–6. (4) Aneurocoris insolitus, (5) Ctenipocoris africanus, (6) Naucoris obscuratus.

Figures 7–9. Species of Laccocoris. (7) Laccocoris limicola, (8) Laccocoris limigenus, (9) Laccocoris spurcus congoensis.

Figures 10–16. Species of Neomacrocoris. (10) Neomacrocoris bondelaufa, (11) Neomacrocoris handlirschi, (12) Neomacrocoris karimii, (13) Neomacrocoris parviceps ocellatus, (14) Neomacrocoris parviceps parviceps, (15) Neomacrocoris usambaricus, (16) Neomacrocoris vuga. 60

Figures 17–18. Species of Macrocoris. (17) Macrocoris flavicollis, (18) Macrocoris sp. (undescribed).

Figures 19–20. Subgenital plates. (19) Neomacrocoris usambaricus, (20) Neomacrocoris vuga.

Figures 21–24. Male genitalia. (21) Neomacrocoris bondelaufa, (22) Neomacrocoris parviceps, (23) Neomacrocoris karimii, (24) Neomacrocoris vuga.

Figures 25–26. Lateral view of pronotum. (25) Neomacrocoris handlirschi, (26) Neomacrocoris parviceps.

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APPENDIX

List of collection sites: 2010, 2011, 2012.

L-1144 TANZANIA : Mwanza Region, Kisesa, ca. 20 km NE Mwanza, 02º33.345'S 33º03.652'E, 1230 m; 24 July 2010; L-1144 colls: R. W. Sites & A. Mbogho, vegetated pond margins

L-1145 TANZANIA: Mwanza Region, Kisesa, ca. 20 km NE Mwanza 02º33.266'S 33º03.767'E 24 July 2010; 1222 m; L-1145 colls: R. W. Sites & A. Mbogho vegetation in pooled cattle stream

L-1146 TANZANIA: Mara Region Tairo, ca. 5 km E of Bunda 02º33.827'S 33º45.695' E 1230 m; 24 July 2010; L- 1146 colls: R. W. Sites & A. Mbogho, heavily vegetated pond margins

L-1147 TANZANIA: Mara Region Ushashi, ca. 5 km N of Nyiendo 02º00.540'S 33º55.241'E 1344m; 24 July 2010; L-1147 colls: R. W. Sites & A. Mbogho. Vegetated pond margins

L-1148 TANZANIA: Mara Region Tamau; 1138 m, 02º06.083'S 33º52.070'E 24 July 2010; L-1148 colls: R. W. Sites & A. Mbogho. shallow vegetated cattle pond

L-1149 TANZANIA: Mwanza Region, near Kashishi; 1344 m, 02º16.239'S 033º48.314'E 24 July 2010; L-1149 colls: R. W. Sites & A. Mbogho vegetated pond margins

L-1150 TANZANIA: Mwanza Region Ihimbili; 1142 m 02º35.247'S 33º24.859'E 24 July 2010; L-1150 colls: R. W. Sites & A. Mbogho, shallow wetland

L-1151 TANZANIA: Mwanza Region Nyanguge; 1159 m 02º32.619'S 33º13.181'E 24 July 2010; L-1151 colls: R. W. Sites & A. Mbogho, large vegetated pond, no cattle usage

L-1152 TANZANIA: Mwanza Region Mwanza, Kirumba, Lake Victoria, 02º30.814'S 33º53.898'E, 1140 m; 25 July 2010 ; L-1152 colls: R. W. Sites & A. Mbogho, low-lying lake margin w/ heavy vegetation

L-1153 TANZANIA: Mwanza Region, 2 km E of Ibanda; 1159 m, 02º46.773'S 33º29.772'E, 25 July 2010, L-1153, colls: R. W. Sites & A. Mbogho, vegetated roadside ditch, with cattle usage

L-1154 TANZANIA: Kagera Region ca. 5 km E of Bhiaramulo, 02º38.159'S 31º20.789'E, 1427 m; 26 July 2010; L-1154 colls: R. W. Sites & A. Mbogho roadside pool with rasses, algae, lily pads

L-1156 TANZANIA: Kagera Region Benako Reservoir; 1492m 02º28.081’S 30º51.355'E, 26 July 2010; L-1156 colls: R. W. Sites & A. Mbogho, marginal vegetation on dam

L-1157 TANZANIA: Kagera Region Karenge; 1234 m 03º07.936'S 31º04.530'E 27 July 2010; L-1157 colls: R. W. Sites & A. Mbogho, shallow pools with heavy, vegetation and cattle use

L-1158 TANZANIA: Kigoma/ Kagera Region border Muvozi River; 1156 m 03º11.331'S 31º02.129'E 27 July 2010; L-1158 colls: R. W. Sites & A. Mbogho, rocky stream with dense filamentous vegetn

L-1159 TANZANIA: Kigoma Region Kiziguzigu stream; 1197 m 03º18.491'S 30º56.207'E, 27 July 2010; L-1159, colls: R. W. Sites & A. Mbogho, small shaded forest stream

L-1160 TANZANIA: Kigoma Region, W of Kibondo Mpemvyi River, 03º44.028'S 30º41.945'E 122 m; 27 July 2010 ; L-1160, colls: R. W. Sites & A. Mbogho, pooled areas of slow stream

L-1161 TANZANIA: Kigoma Region Nyakivaya Lake, ca. 5 km N of Kasulu; 1165 m 04º16.455'S 30º26.111'E, 27 July 2010 ; L-1161,colls: R. W. Sites & A. Mbogho, shallow small lake with heavy vegetation

L-1162 TANZANIA: Kigoma Region, Luchugi River, 1217 m, 04º31.733'S 30º09.699'E, 27 July 2010, L- 1162,colls: R. W. Sites & A. Mbogho, small stream

L-1163 TANZANIA: Kigoma Region, tributary to Kalembela River, 04º40.743'S 30º03.994'E 1244 m; 27 July 2010; L-1163, colls: R. W. Sites & A. Mbogho, irrigation canal

L-1164 TANZANIA: Kigoma Region Kalembela River, 1249 m, 04º40.813'S 30º03.964'E, 27 July 2010, L-1164, colls: R. W. Sites & A. Mbogho, marginal vegetation in slow current

L-1165 TANZANIA: Kigoma Region, 40 km N of Kigoma, 952 m, 04º53.201'S 29º52.190'E 27 July 2010, L-1165 colls: R. W. Sites & A. Mbogho, rapid stream w/ slab rock, gravel, marginal vegetation

L-1166 TANZANIA: Kigoma Region, Lake Tanganyika, Kigoma, 04º54.719'S 29º37.989'E, 772 m; 28 July 2010; L-1166, colls: R. W. Sites & A. Mbogho, rootmats of matete on waveswept sandy shoreline

L-1167 TANZANIA: Kigoma Region Lake Tanganyika, Kigoma, 04º55.319'S 29º40.256'E, 773 m; 28 July 2010; L-1167 colls: R. W. Sites & A. Mbogho, dense rootmats of matete in heavy wave action

L-1168 TANZANIA: Kigoma Region 5 km N of Kigoma stream inlet to Lake Tanganyika, 04º51.799'S 29º37.786'E,770 m ; 28 July 2010 ; L-1168, colls: R. W. Sites & A. Mbogho, emergent grasses in standing water 71

L-1169 TANZANIA: Kigoma Region, 15 km N of Kigoma Mungonya River; 04º50.337'S 9º40.001'E 806 m; 28 July 2010; L-1169, colls: R. W. Sites & A. Mbogho, muddy bank with sparse marginal vegetation

L-1170 TANZANIA: Kigoma Region Msimba, ca. 18 km N of Kigoma, 04º51.058'S 29º42.552'E, 807 m, 28 July 2010; L-1170, colls: R. W. Sites & A. Mbogho, very slowly flowing stream with marginal vegetation

L-1171 TANZANIA: Kigoma Region, Katosho Reservoir, 8 km E of Kigoma, 802 m, 04º51.221'S 29º39.436'E, 28 July 2010, L-1171 colls: R. W. Sites & A. Mbogho, emergent dead tall grasses in turbid reservoir

L-1172 TANZANIA: Kigoma Region, Lake Tanganyika, Kigoma, Ujiji, 04º54.941'S 29º36.661'E, 763 m; 28 July 2010; L-1172, colls: R. W. Sites & A. Mbogho, cobble substrate with submerged veg and wave action

L-1173 TANZANIA: Kigoma Region, Gombe National Park; Mwagongo, Nkonya River, 4º37.351'S 29º38.319'E, 775 m; 29 July 2010; L-1173, colls: R. W. Sites & A. Mbogho, ravel/ rocky clear stream with sparse marginal veg

L-1174 TANZANIA: Kigoma Region Gombe National Park; Mitumba stream; 776 m, 04º38.405'S 29º37.823'E, 29 July 2010; L-1174, colls: R. W. Sites & A. Mbogho, heavily vegetated narrow forest stream

L-1175 TANZANIA: Kigoma Region, Gombe National Park; Rutanga stream; 776 m, 04º39.111'S 29º37.660'E, 29 July 2010; L-1175, colls: R. W. Sites & A. Mbogho, sandy & muddy stream w/ heavy vegetation

L-1176 TANZANIA: Kigoma Region, Gombe National Park, Kakombe Waterfall, 29 July 2010; L-1176 colls: R. W. Sites & A. Mbogho, wet rocks sprayed by falls

L-1177 TANZANIA: Kigoma Region Gombe National Park; stream from Kakombe waterfall at Lake Tanganyika; 1177 m, 04º40.197'S 29º37.376'E, 29 July 2010 ; L-1177 colls: R. W. Sites & A. Mbogho rocky stream with little vegetation

L-1178 TANZANIA: Kigoma Region Gombe National Park, Mukenke stream ; 770m 04º40.662'S 29º37.246'E 29 July 2010 ; L-1178 colls: R. W. Sites & A. Mbogho small rocky stream w/o vegetation

L-1179 TANZANIA: Kigoma Region Gombe National Park; 767 m 04º42.002'S 29º38.856'E 29 July 2010 ; L-1179 colls: R. W. Sites & A. Mbogho, algae covered cobble of riffle & muddy pool

L-1180 TANZANIA: Kigoma Region Gombe National Park ; Elev.769 m 04º42.264’S 29º36.802'E 29 July 2010 ; L-1180 colls: R. W. Sites & A. Mbogho, rocky riffles with minimal algae

L-1181 72

TANZANIA: Kigoma Region Gombe National Park ; Buave stream; 766 m, 04º43.949'S 29º36.560'E, 29 July 2010 ; L-1181,colls: R. W. Sites & A. Mbogho, small rocks, marginal rootmats and vegetation

L-1182 TANZANIA: Kigoma Region Kazuramimba ; 1082 m 05º00.155'S 30º00.971'E 30 July 2010 ; L- 1182 colls: R. W. Sites & A. Mbogho, muddy pools of small stream with marginal vegetation

L-1183 TANZANIA: Kigoma Region Kibaoni stream ; 1026 m 05º03.974'S 30º20.666'E 30 July 2010 ; L- 1183 colls: R. W. Sites & A. Mbogho, pooled areas of slow stream

L-1184 TANZANIA: Rukwa Region, 05º12.863'S 30º23.357'E, 1206 m; 30 July 2010; L-1184, colls: R. W. Sites & A. Mbogho, slow stream pools w/ algae and overhanging vegetation

L-1185 TANZANIA: Rukwa Region, 110 km N of Mpanda; 1295 m, 05º37.745'S 30º36.039'E 30 July 2010; L- 1185 colls: R. W. Sites & A. Mbogho, shallow creek w/ muddy bottom and marginal grasses

L-1186 TANZANIA: Rukwa Region, 93 km N of Mpanda; Kapipula stream; 1326 m, 05º42.233'S 30º41.563'E, 30 July 2010; L-1186, colls: R. W. Sites & A. Mbogho, stream wetland w/ dense emergent grasses

L-1187 TANZANIA: Rukwa Region, 57 km N of Mpanda, Masito/Ogala, 1144 m, 05º56.742'S 30º57.812'E, 30 July 2010, L-1187, colls: R. W. Sites & A. Mbogho, slow stream, coarse roots at margin

L-1188 TANZANIA: Rukwa Region, Mpanda stream, Mpandandogo, 06º12.124'S 30º59.284'E, 1146 m; 30 July 2010; L-1188, colls: R. W. Sites & A. Mbogho, slow stream, overhanging grasses

L-1189 TANZANIA: Rukwa Region Lake Tanganyika, Karema, 06º49.694'S 30º26.646'E, 780 m; 31 July 2010; L- 1189 colls: R. W. Sites & A. Mbogho, heavily wave swept roots of reed grass

L-1190 TANZANIA: Rukwa Region, 1 km E of Karema; 782 m, 06º48.473'S 30º26.315'E, 31 July 2010; L-1190, colls: R. W. Sites & A. Mbogho, algal mat at margin

L-1191 TANZANIA: Rukwa Region 35 km E of Karema; 951 m, 06º43.726'S 30º33.681'E, 31 July 2010 ; L-1191 colls: R. W. Sites & A. Mbogho, slow stream, big pools and algal slick

L-1192 TANZANIA:Rukwa Region 39 km E of Karema; 966m, 06º42.830'S 30º34.453'E 31 July 2010 ; L-1192 colls: R. W. Sites & A. Mbogho, bedrock, connected pools, heavy filamentous algae

L-1193 TANZANIA: Rukwa Region, ca. 54 km E. of Karema, 1086 m; 31 July 2010; L-1193, 06º37.275'S 30º39.593'E colls: R. W. Sites & A. Mbogho, shaded stream, negligible flow 73

L-1194 TANZANIA: Rukwa Region, ca. 64 km E Karema; 1115 m 06º32.757'S 30º41.956'E, 31 July 2010 L- 1194 colls: R. W. Sites & A. Mbogho, muddy cattle pool w/ emergent grasses

L-1195 TANZANIA: Rukwa Region, Mpanda, Kasimba River, 06º21.602'S 31º04.382'E, 1064 m; 1 August 2010; L-1195 colls: R. W. Sites & A. Mbogho, grasses hanging into river

L-1196 TANZANIA: Rukwa Region, Katavi National Park; Lake Katavi; 961 m, 06º41.882'S 31º01.376'E,1 August 2010 ; L-1196 colls: R. W. Sites & A. Mbogho, grass clumps in muddy seasonal wetland

L-1197 TANZANIA: Rukwa Region, Katavi National Park, Katuma River wetland; 965 m, 06º39.307'S 31º05.040'E, 1 August 2010, L-1197 colls: R. W. Sites & A. Mbogho, vegetated shallow wetland

L-1198 TANZANIA: Rukwa Region, Katavi National Park , spring at Ikuu ; 915 m , 06º53.090'S 31º11.204'E , 1 August 2010 ; L-1198, colls: R. W. Sites & A. Mbogho, helocrene with hippos

L-1199 TANZANIA: Rukwa Region, Mbede River at Mbede 07º12.546'S 31º18.861'E 951 m; 1 August 2010; L-1199 colls: R. W. Sites & A. Mbogho gravel silt substrate w/overhanging vegetation

L-1200 TANZANIA: Rukwa Region Sau River at Kasunga; 1795 m 08º03.444'S 31º30.112'E, 2 August 2010 ; L-1200 colls: R. W. Sites & A. Mbogho , marginal dense vegetation in cattle pool of stream

L-1201 TANZANIA: Rukwa Region 20 km SW of Subawanga ; Katuka River at Katuka 08º06.641'S 31º31.786'E 1858 m; 2 August 2010; L-1201; colls: R. W. Sites & A. Mbogho pools of stream and narrow channel w/ dense vegetation

L-1202 TANZANIA: Rukwa Region, Msanzi River at Msanzi, 08º11.411'S 31º31.295'E, 1778 m; 2 August 2010; L-1202; colls: R. W. Sites & A. Mbogho, rocks, mud, gravel, sand, marginal grasses

L-1203 TANZANIA: Rukwa Region, Lake Tanganyika; Kasanga, 08º26.746'S 31º08.642'E, 773 m; 2 August 2010 ; L-1203 colls: R. W. Sites & A. Mbogho, waveswept reed grass clumps

L-1204 TANZANIA: Rukwa Region, Galasia River at Kayengesa, 08º15.146'S 31º42.284'E, 1682 m; 3 August 2010; L-1204 colls: R. W. Sites & A. Mbogho, mud substrate w/ little vegetation

L-1205 TANZANIA: Rukwa Region, Kamawe River at Puwi, 08º21.029'S 31º50.054'E, 1598 m; 3 August 2010; L-1205 colls: R. W. Sites & A. Mbogho, clear stream, marginal vegetation, some rocks

L-1206

TANZANIA: Mbeya Region, 1 km N of Nyenjele,Muwawa River, 1448 m, 08º55.289'S 32º21.583'E, 3 August 2010; L-1206; colls: R. W. Sites & A. Mbogho, pool with overhanging grasses

L-1207 TANZANIA: Mbeya Region, Maboo stream at Mbeya Peak village, 2433 m, 08º50.432'S 33º25.060'E, 4 August 2010; L-1207; colls: R. W. Sites & A. Mbogho, vegetated pool of small high elevation stream

L-1208 TANZANIA: Mbeya Region Itunga inlet,Lake Nyassa, 482 m, 09º35.094'S 033º56.769'E, 4 August 2010; L-1208, coll: R. W. Sites and A. Mbogho, Heavy marginal vegetation at boat ramp

L-1209 TANZANIA: Mbeya Region, Lake Nyassa at Itunga; 480m, 09º34.684'S 33º56.881'E, 4 August 2010 ; L- 1209; colls: R. W. Sites & A. Mbogho, root tangles of matete

L-1210 TANZANIA: Ruvuma Region ca. 5 km N of Songea, Luwila River, 1086 m, 10º37.510'S 35º39.199'E, 5 August 2010 ; L-1210; colls: R. W. Sites & A. Mbogho slow sandy stream w/ overhanging veg

L-1211 TANZANIA: Ruvuma Region 20 km N of Songea; Lumecha River at Samangula ; 994 m, 10º30.149'S 35º40.268'E, 5 August 2010 ; L-1211; colls: R. W. Sites & A. Mbogho overhanging grasses

L-1212 TANZANIA: Ruvuma Region Luhimba River ; 996 m 0º26.742'S 35º40.258'E 5 August 2010 ; L- 1212 colls: R. W. Sites & A. Mbogho overhanging veg, & side pool w/ algae and veg

L-1213 TANZANIA: Ruvuma Region Hanga River at Ngadinda 10º10.264'S 35º39.611'E 777 ; 5 August 2010 ; L-1213 colls: R. W. Sites & A. Mbogho overhanging veg and matete

L-1214 TANZANIA: Iringa Region, nr Nyololo,1821 m, 08º25.911'S 35º11.684'E, 6 August 2010 ; L-1214; colls: R. W. Sites & A. Mbogho grassy seasonal wetland

L-1215 TANZANIA: Iringa Region Mwiriwa River at Kimamba 07º39.657'S 36º05.275'E 1312 m; 6 August 2010; L-1215 colls: R. W. Sites & A. Mbogho pooled stream w/ algae and overhanging grasses

L-1216 TANZANIA: Morogoro Region Ruaha River at Mbuyuni; 527 m, 07º27.922'S 36º30.973'E 6 August 2010; L-1216 colls: R. W. Sites & A. Mbogho, large river w/ sparse marginal vegetation

L-1218 TANZANIA: Morogoro Region, Mikumi National Park; 520 m, 07º20.432'S037º07.048'E 7 August 2010; L-1218 colls: R. W. Sites & A. Mbogho, muddy hippo pond

L-1219 TANZANIA: Morogoro Region Mikumi National Park ; 529 m 07º21.608'S 37º03.337'E 7 August 2010 ; L-1219 colls: R. W. Sites & A. Mbogho heavily vegetated seasonal pond

L-1220 TANZANIA: Morogoro Region ca. 21 miles NW of Morogoro, 06º29.634'S 37º33.465'E, 372 m; 7 August 2010; L-1220 colls: R. W. Sites & A. Mbogho, heavily vegetated pond ringed with reed grass

L-1221 TANZANIA: Morogoro Region, 40 miles NE of Morogoro,06º24.257'S 37º25.467'E,401 m; 7 August 2010; L-1221 colls: R. W. Sites & A. Mbogho, Shallow cattle pond, sparse vegetation

L-1222 TANZANIA: Morogoro Region, Magole River at Dumila; 436 m, 06º23.221'S 37º21.629'E 7 August 2010 ; L-1222 colls: R. W. Sites & A. Mbogho marginal veg in shallow sandy river

L-1223 TANZANIA: Dodoma Region, Lukali Hot spring, 991m, 05º43.936'S 35º30.082'E, 8 August 2010 ; L- 1223; colls: R. W. Sites & A. Mbogho warm spring with algae and Typha angustifolia

L-1224 TANZANIA: Iringa Region, Silunga River at Chikuyu, 05º52.223'S 35º04.568'E, 842 m; 8 August 2010; L-1224 colls: R. W. Sites & A. Mbogho, dead twigs in muddy river

L-1225 TANZANIA: Iringa Region, Kipumbuiko River, 1516m, 05º05.667'S 34º45.940'E 8 August 2010; L-1225; colls: R. W. Sites & A. Mbogho, shallow silty stream w/ algae & Typha angustifolia

L-1226 TANZANIA: Singida Region, Kindai Reservoir at Singida, 04º50.065'S 34º44.173'E, 1484 m; 8 August 2010; L-1226; colls: R. W. Sites & A. Mbogho, Tall grasses

L-1227 TANZANIA: Singida Region, Singidan Reservoir at Singida, 04º47.568'S 34º44.570'E 1483 m; 8 August 2010; L-1227; colls: R. W. Sites & A. Mbogho, Typha angustifolia roots & submergent vegetation

L-1228 TANZANIA: Singida Region, Ifombou; 1546 m, 04º40.323'S 34º50.591'E 9 August 2010; L-1228, colls: R. W. Sites & A. Mbogho, seasonal cattle pond w/ Typha angustifolia and submergent vegetatn

L-1229 TANZANIA: Singida Region, Mtinko stream at Mtinko, 04º33.525'S 34º50.649'E, 1552 m; 9 August 2010; L-1229 colls: R. W. Sites & A. Mbogho, pooled temp stream w/ algae

L-1230 TANZANIA: Singida Region, Mwanga River at Mwanga, 04º17.496'S 34º51.969'E, 1531 m; 9 August 2010; L-1230 colls: R. W. Sites & A. Mbogho, stream pools w/ algae and marginal vegetation

L-1231 TANZANIA: Manyara Region, Ghoro ng'aida River at Haydom, 04º11.487'S 35º01.878'E 1681 m; 9 August 2010; L-1231; colls: R. W. Sites & A. Mbogho, pooled discontinuous stream

L-1232 TANZANIA: Manyara Region, Yayeda River at Bagheti, 04º00.158'S 35º20.470'E, 1974; 9 August 2010; L-1232 colls: R. W. Sites & A. Mbogho, marginal overhanging grasses

L-1233 TANZANIA: Manyara Region, Tiawi ; 1951 m 03º54.742'S 35º28.008'E 9 August 2010 ; L-1233 colls: R. W. Sites & A. Mbogho temporary vegetated pond

L-1234 TANZANIA: Manyara Region, Tiawi Kwaanseri Lake, 03º54.600'S 35º27.739'E, 1940 m; 9 August 2010; L-1234 colls: R. W. Sites & A. Mbogho wetland with heavy vegetation

L-1235 TANZANIA: Arusha Region, Kirurum River at N end of Manyara National Park, 03º22.094'S 35º50.418'E,1012 m; 10 August 2010; L-1235 colls: R. W. Sites & A. Mbogho,rocky high velocity stream

L-1236 TANZANIA: Arusha Region, Simba stream ; 973 m, 03º22.399'S 35º51.114'E, 11 August 2010 ; L- 1236; colls: R. W. Sites & A. Mbogho irrigation canal w/ overhanging grasses

L-1237 TANZANIA: Arusha Region, Nadosaito Reservoir, Rokado (Maasai village) ; 1313 m, 03º25.137'S 36º22.880'E 11 August 2010 ; L-1237 colls: R. W. Sites & A. Mbogho, vegetated seasonal lake

L-1238 TANZANIA: Arusha Region, 0.5 km S of Kisongo; 1369 m 03º22.853'S 36º33.552'E 11 August 2010 ; L-1238 colls: R. W. Sites & A. Mbogho temporary pond with grasses

L-1239 TANZANIA: Arusha Region, Sanaware River at Arusha, 03º21.549'S 36º41.619'E, 1442 m; 11 August 2010; L-1239; colls: R. W. Sites & A. Mbogho, shaded rocky creek below elementary school

L-1240 TANZANIA: Arusha Region, Temi River at Arusha; 1444 m, 03º21.824'S 36º42.173'E 11 August 2010 ; L-1240 colls: R. W. Sites & A. Mbogho, shallow, gravel/pebble bottom, marginal veg

L-1241 TANZANIA: Kilimanjaro Region, Hona River at Himo ; 836 m, 03º23.416'S 37º32.689'E 11 August 2010; L-1241 colls: R. W. Sites & A. Mbogho, clear, rocky stream w/ emergent vegetation

L-1242 TANZANIA: Kilimanjaro Region, Mauna; 1631 m, 03º16.765'S 37º28.453'E 12 August 2010; L-1242 colls: R. W. Sites & A. Mbogho small grassy channel at District office

L-1243 TANZANIA: Kilimanjaro Region Mae stream at Rua village 03º15.053'S 37º27.757'E, 1860 m; 12 August 2010; L-1243 colls: R. W. Sites & A. Mbogho, grassy high elevation stream

L-1244 TANZANIA Kilimanjaro Region, Mae Waterfall at Rua village 03º15.079'S 37º28.152'E 1803 m; 12 August 2010; L-1244 colls: R. W. Sites & A. Mbogho waterfall w/ veg at perimeter of plunge pool

L-1245 TANZANIA: Kilimanjaro Region Mae River at Mauna ; 1660 m 03º16.65'S 37º28.29'E 12 August 2010 ; L-1245 colls: R. W. Sites & A. Mbogho mountain rock/gravel stream

L-1246 TANZANIA: Kilimanjaro Region roadside waterfall 1442 m 03º17.439'S 37º29.854'E 12 August 2010 ; L-1246 colls: R. W. Sites & A. Mbogho algae and moss on waterfall

L-1247 TANZANIA: Kilimanjaro Region Marwa ; 647 m 04º06.018'S 37º32.009'E 13 August 2010 ; L-1247 colls: R. W. Sites & A. Mbogho, grassy ditch outflow from Ruvu River

L-1248 TANZANIA: Tanga Region, Mkomazi; 465 m, 04º38.685'S, 38º04.037'E 13 August 2010 ; L-1248 colls: R. W. Sites & A. Mbogho temporary flooded area from Mkomazi River

L-1249 TANZANIA: Tanga Region, Pangani River at Bwiko; 534 m, 04º39.630'S 38º02.105'E, 13 August 2010; L-1249, colls: R. W. Sites & A. Mbogho, margins of large river

L-1250 TANZANIA: Kilimanjaro Region, Saseni River, 520 m, 04º28.315'S 38º04.665'E, 13 August 2010; L- 1250; colls: R. W. Sites & A. Mbogho, shallow sandy river w/ submerged vegetation

L-1251 TANZANIA: Kilimanjaro Region, Jiwemawe Reservoir at Ndungu, 04º25.496'S 38º05.201'E 515 m; 13 August 2010; L-1251; colls: R. W. Sites & A. Mbogho, vegetated margins of reservoir & below spillway

L-1252 TANZANIA: Tanga Region Mazinde stream at Mazinde 04º48.209'S 38º12.615'E 433 m; 13 August 2010; L-1252 colls: R. W. Sites & A. Mbogho narrow stream with tall emergent macrophytes

L-1253 TANZANIA: Tanga Region, river from Soni to Mombo, 2 km E of Vuga Junction, 04º52.040'S 38º20.906'E, 460 m; 14 August 2010; L-1253 colls: R. W. Sites & A. Mbogho, grassy overhangs in rocky streams

L-1254 TANZANIA: Tanga Region, river from Soni to Mombo, Karense, 1 km E of Soni, 04º50.284'S 38º22.355'E, 1051 m; 14 August 2010; L-1254, colls: R. W. Sites & A. Mbogho, vegetated overhangs and undercuts

L-1255 TANZANIA: Tanga Region, Gomba River, 2 km S of Chekelei, 04º50.284'S 38º18.774'E 412 m; 14 August 2010; L-1255; colls: R. W. Sites & A. Mbogho, grassy overhangs in irrigation canal

L-1256 TANZANIA: Tanga Region, Kwamaluli River at Mbagai, 04º58.561'S 38º23.154'E, 465 m; 14 August 2010; L-1256 colls: R. W. Sites & A. Mbogho, shaded narrow stream w/ overhanging veg and undercuts

L-1257 TANZANIA: Tanga Region, Korogwe; 301 m, 05º09.165'S 38º26.688'E, 14 August 2010; -1257; colls: R. W. Sites & A. Mbogho, muddy temporary stream

L-1258 78

TANZANIA: Tanga Region Kwamkuyo River Karimi Tea Estates; 901 m, 05º04.491'S 38º38.215'E, 15 August 2010; L-1258 colls: R. W. Sites & A. Mbogho, sandy/silty stream w/ cultivated overhanging veg

L-1259 TANZANIA: Tanga Region Derema River, Karimi Tea Estates; 809 m, 05º04.895'S 38º38.548'E 15 August 2010; L-1259; colls: R. W. Sites & A. Mbogho, veg margins w/ heavy red siltation

L-1260 TANZANIA: Tanga Region, Kihuuwi River at Timba; 174 m, 05º07.683'S 38º41.418'E 15 August 2010; L- 1260 colls: R. W. Sites & A. Mbogho, marginal grasses with silty/rocky bottom

L-1261 TANZANIA: Tanga Region Jembe stream at Muheza; 182 m, 05º10.260'S 38º47.106'E, 15 August 2010; L-1261; colls: R. W. Sites & A. Mbogho, grassy overhangs in urban stream

L-1262 TANZANIA: Pwani Region Wami River at Wami Bridge, 06º14.876'S 38º23.230'E, 93 m; 16 August 2010; L-1262; colls: R. W. Sites & A. Mbogho, large muddy river with grasses and matete

L-1263 TANZANIA: Pawni Region, pond by Ruvu River W of Bagamoyu, 2 m 06º28.545'S 38º50.614'E 16 August 2010; L-1263 colls: R. W. Sites & A. Mbogho,water lettuce and other vegetation

L-1264 TANZANIA: Zanzibar, Mjinimagharibi Region, Bumbwisudi; 41 m,06º03.526'S 39º15.680'E 17 August 2010 ; L-1264 colls: R. W. Sites & A. Mbogho, temporary roadside ditch

L-1265 TANZANIA: Zanzibar, Kaskazini Region, Kitope River at Zingwezingwe Bridge, 06º00.963'S 39º14.734'E, 39 m; 17 August 2010; L-1265 colls: R. W. Sites & A. Mbogho, discontinuous pools w/ aquatic veg and grasses

L-1266 TANZANIA: Zanzibar, Kaskazini Region, Kitope River at Mkaratini Bridge, 06º00.714'S 39º14.841'E, 30 m; 17 August 2010; L-1266; colls: R. W. Sites & A. Mbogho, discontinuous pools w/ aquatic veg and grasses

L-1267 TANZANIA: Zanzibar Kusini Region, Mwera River at Mwera, 06º08.811'S 39º16.264'E 23 m; 17 August 2010; L-1267, colls: R. W. Sites & A. Mbogho, flooded, vegetated field margin

L-1322 TANZANIA: Tanga Region on Usambara Mountains, at River from Soni to Mombo, 3.5 km E of Vuga Junction, 04º52.471'S 038º20.660'E, 992 m; 28 December 2011; L-1323 colls: A. Y. Mbogho. Overhang grass, rocky flowing stream

L-1323 TANZANIA: Tanga Region, on Usambara Mountains, River from Soni to Mombo, at Mawei, 1.5 km E of Soni, 04º50.527'S 038º22.238'E 1184 m; 28 December 2011; L-1323; colls: A. Y. Mbogho. Vegetated overhangs/undercuts/overflowg

L-1324 TANZANIA: Morogoro Region, Uldizungwa National Park, Ruaha River at Iyovi; 477 m, 07º34.873'S 036º47.494'E 30 December 2011; L-1324; colls: A.Y. Mbogho. A Large river with large marginal vegetation

L-1332 TANZANIA: Tanga Region on Usambara Mountains, at River from Soni to Mombo 3.5 m E of Vuga Junction 04º52.471'S 038º20.660'E 992 m; 01 February, 2012; L-1323 colls: A. Y. Mbogho. Overhang grass, rocky flowing stream

L-1333 TANZANIA: Tanga Region on Usambara Mountains, River from Soni to Mombo, at Mawei, 1.5 km E of Soni 04º50.527'S 038º22.238'E 1184 m; 01 February, 2012; L-1323; colls: A. Y. Mbogho. Vegetated overhangs/undercuts/overflowg

L-1334 TANZANIA: Morogoro Region, Uldizungwa National Park, Ruaha River at Iyovi; 477 m 07º34.873'S 036º47.494'E 29 January 2012; L-1324 colls: A.Y. Mbogho. A Large river with large marginal vegetation

VITA

Aaron Y. Mbogho was born on November 27, 1969 in Singida Region and raised at

Mtinko village in the central part of Tanzania. He received both primary and secondary school education in Singida: at Malolo Primary School (1986) and Mwenge Secondary

School (1990 and 1993) respectively. He received his BSc degree in Horticulture with honors from Sokoine University of Agriculture (SUA), Morogoro Tanzania in 2002. He then earned his MSc degree in Entomology from the University of Missouri (MU),

Columbia, USA in 2012. He also completed a course on Sustainable Agriculture at

Navdanaya Agro-Ecological and Biodiversity Conservation Centre in Dehradun, India.

Mr. Mbogho is a Tanzanian civil servant employed by the Ministry of

Agriculture, Food and Cooperatives as Agro-Officer responsible for Plant Health

Services. During his first research work in Diamondback Moth control in 2002, he was guided by Dr. Myron Zalucki, professor of Zoology and Entomology at University of

Queensland, Australia and he is the first person to work on the Economic Threshold for

Diamondback Moth in Cabbage in Tanzania. He is an apprentice of Dr. Ian F. Grant, a professor of Ecotoxicology at the Natural Resources Group (the NR-Group) UK and he is the first person to conduct Rapid Risk Assessment (RRA) of Pesticides and he may still be the only person with such RRA capability in Tanzania. He has co-authored several papers with researchers from SUA and MU. He contributed to the first two patents from

SUA: the Portable Double Sided Steel Frame for Log Sawing (Patent # TZ/P/07/00140) and the Tree Pusher (Patent # TZ/P/07/00143). While pursuing his master’s degree at MU

81 under the advisory and supervision of Dr. Robert W. Sites, a professor of Entomology, he developed a taxonomic key for 14 known species of Naucorids of Tanzania to include the two new species: Neomacrocoris bondelaufa and Neomacrocoris vuga they described in

2012. He is a recipient of Thurgood Marshall Fellowship, the prestigious academic award made by Graduate School. Amongst numerous national and international awards and recognitions embracing his rich experience, it worth mentioning that he won first place in Student Competition, Entomological Society of America President’s Prize in

Systematics, Evolution, and Biodiversty category in 2010. He is a permanent member of the Golden Key International Honour Society validated by University of Missouri,

Columbia for his academic excellence. As a plant protectionist with a precision agriculture vision, he believes and advocates for Integrated Pest Management techniques.

He enjoys the most when working in Field Crop Entomology especially in the philosophical ventures such as Economic Thresholds for insect pests.