^ ^ kJ & m b er1 0 I S & - F 7 C tf2 (a * 7 1 2 6 $ ) University of Ghana http://ugspace.ug.edu.gh the distribution, westing habits, and activity pattern OP CAMPONOTUS ACVAPIMENSIS MAYR (rIYI IEt'IOPTERA PORI'UCIDAE) IN SAVANNA AREAS OF SOUTHERN GHANA. By KOFI OSAE OFEI B.Sc. (Ed)., B. Sc. (Hons) A thesis submitted for the Master of Science degree at the University of Ghana, 1978. University of Ghana http://ugspace.ug.edu.gh ZOOLOGY D3PARMOT, UNIY3HSITT OF GEA1TA LEGON This is to certify that this thesis has not been submitted for a degree to any other University. t is entirely my own work, and all help has been dvly acknowledged. KOFI 03AS 0F3I University of Ghana http://ugspace.ug.edu.gh This w©rk is dedicated t© my late father who through his needy days sacrificed and made me what I am today. His memories will always remain* University of Ghana http://ugspace.ug.edu.gh ill ACKNOWLEDGEMMT This w®rk is a report of 1 2 months of field and laboratory studies# My gratitude goes to Professor R. Kumar, my immediate supervisor, who suggested this project and without whose encourage­ ment, directions and criticisms this work would not have been possible# My thanks go to the members of my supervisory committee especially Dr* Lock of Botany Department University of Ghana whose suggestions and directions highlighted the problems of the study plots and suggested alternative methods to study them. I wish to thank also, the Headmaster of Presbyterian College Legon, Rev. E.S. Mate-Eodjo for permitting me to do the course at a veiy short notice, and allowing me to use the laboratory facilities and portions cf the school plots as my study plots without disturbances. I am also indebted to him for his help which enabled me to enjoy study leave and stay on the school compound during the course* I wish to thank Mr. D. Decker, Chief Technician, Department of Zoology for his help in providing me with various materials at odd hours. I thank Mr Afreh Nuamah, a National Service worker of the Zoology Department who helped me with the operation of the scientific equipments used for field work. University of Ghana http://ugspace.ug.edu.gh I thank Mr. R. Pobi formerly of Presbyterian College for typing various portions of this work for me# X wish, to thank Mr. E. Attua-Afari, the Assiatant Headmaster, who permitted me to use the school typewriter for the initial typing work. I thank Miss Gloria Aye of Cocoa Marketing Board, Produce Buying Section Koforidua for spending her precious hours to type the final work. My thanks are also extended to Mrs. Jackson a Horticulturist, now at Kumasi University for helping me with the identification of the various plant species that were on the study plots. Lastly, my thanks go to the Department of Zoology and Ghana Education Service who supported this work. iv University of Ghana http://ugspace.ug.edu.gh TABLE OP CONTECTTS University of Ghana http://ugspace.ug.edu.gh TABLE OF CONTENTS 1 General Introduction 2 Genus Camponotus Mayr 3 The study area • • • • • * 3.1 Locality ♦♦ 3.2 Plot A (savanna grassland) •* 3.3 Effects of wet season on plot A * • 3.4 Effects of dry season on plot A • ♦ 3*5 Plot B • • • • ♦ • • * 3.6 Effects of wet season on plot B *• •• 3.7 Effects of dry season on plot B •• 3.8 Other disturbances affecting the 2 plots. 4 Relative distribution of ants in space 4.1 Introduction •• •• 4.2 Methods •• •* *• •• 4.2.1 Abundance of nests and nest distribution in relation to shade regimes on plot A. 4.2.2 Abundance of nests and nest distribution in relation to shade regimes on plot B. 4.2.3 Colony size 4.2.4 Distribution of Gamp, acvapimensis in relation to other ants on plot A •♦ •♦ • * 4.2.5 Distribution of Camp, acvapimensis in relation to other ants on plot B. • • •« 4*3 Results / Observations •• •• * • University of Ghana http://ugspace.ug.edu.gh - vii - 4.3.1 Abundance of nests on plot A. • • « . page 25 4.3.2 Abundance of nests on plot B. . * • * 24 4.3.3 Distribution in relation to shade regimes 25 4.3.3.1 On plot A . • .. • • • • 25 4.3.3.2 On plot B . * . • • • • • 20 4.3.4 Colony size •» •. • • • • 30 4.4 Discussion .* *. • • ♦ • •2- A 4.4.1 Abundance of nests on plot A • • • • 34 4.4.2 Abundance of nests on plot 3 * • • * 3 G 4.4.3 Distribution of the nests •» • • Jv 4.4.3*1 Colony size .» .. • • « • 36 4*4.4 Distribution in relation to other ants 36 4.4.5 Nests and nesting habits •0 0 « • 0 38 4.4.5.1 Camp, acvapimensis and Camp. seric«us 3S 4.4.5.1.1 Numerical abundance 0 © « « so 4.4.5.2 Camp* acvapimensis and Camp, vestitus 40 4.4.5.3 Camp, acvapimensis and Annoma • • • • 4C 4.4.5 *3.1 Numerical abundance • * • « • t 41 4.4.5.4 Camp, acvapimensis and Platythyreus tarsartus 41 5 Nests and nesting habits • * • • 43 5.1 Introduction • • • . • • » • 43 5.? Methods • • « . • 6 45 5.2.1 Sampling proceedure • • «. * 46 5.2.3 Sorting • • • fr » • 47 University of Ghana http://ugspace.ug.edu.gh - Till - Vage 4.8 5*2»3 Counting »• •• 00 •• ^ 5*2.4 3o&d measurements 48 5«2»5 Ehe' nost • • • * * * 0 •• 49 5o2*6 DJrpft ©f soil in nhich nesting takes place 50 5o2*7 Position: of nest entrance in relation to vegetation 5t 5.2o8 Effects of flooding on ants 52 5«2o9 $jrpe of nests .* • • *• •• 5£ 5o3 Observations / Results 52 5P3«t Contents of nests ». *• •• 52 5*3*2 Guests of Campoacvapimensis 56 5*3*3 Phenology of brood and castes, 57 5*3*4 Hejad measurements • • .* • • 59 5*3*5 The nest • * •• •• • • 60 5*3*6 TJype of soil in which nesting takes place 61 5*3*7 Position of nest entrance in relation to vegetation 61 5*3*8 Effects of flooding 011 ants 61 5*3<>9 Nest cleaning *. .. 0. .. 62 5*3*10 Type of nests „• •• 00 • • 62 5*3*11 Arrangements of castes and brood uitliin nests 63 5 . 3*12 Nest abandonment . . * e . . 64 5*4 Discussion ., 09 ., ,. 64 5* 4*1 Contents of nests <,* ,, 64 5* 4*2 Populations in nests and phenology of ants 65 5*4*3 Head measurements ,. .. M 50 5* 4*4 The Nests ## ## .. 6c University of Ghana http://ugspace.ug.edu.gh - ix - page 5.4.5 Soils in which nesting- took place 70 5.4.6 Effects of flooding on nests • • • • 71 5.4.7 Nest cleaning 71 5.4.8 Types of nests and arrangement of caste and brood 71 6 Activity patters of Camponotus acvapimensis 72 6.1 Introduction 72 6.2 Materials and methods • • •• •• 75 6.2.1 Foraging activity 75 6.2.1.1 J Diurnal activity pattern .• •• • • 75 6.2.1.2 I Nocturnal activity pattern •■ •• 76 6.2.2 Foraging space and distance •• • * 77 6.2.3 Induced foraging .. •• •• 73 6.2.4 Ants and phytophagous insects associationship 7S 6.2.5 Flight activity .. •• 79 6.2.6 Food items collected by ants 7°' 6.3 Results / Observations .. • • SO 6.3.1 Foraging activity of Camp.acvapimensis 80 6.3.1.1 Diurnal activity .. • • •• 81 6.3.1.2 Nocturnal activity •. • • •• 84 6.3.2 Induced activity •. 86 6.3.3 Foraging space and distance •• 86 6.3.4 Ants - phytophagous insects associationship 88 6.3.5 Food items collected by ants •* * • 0? 6.3.6 Flight activity .. •• «• ClCO University of Ghana http://ugspace.ug.edu.gh 5.4.5 Soils in which nesting took place «• page 70 5.4.6 Effects of flooding on nests •• ft a 71 5.4.7 Nest cleaning .. •• • • 71 5*4.8 Types of nests and arrangement of caste and rtfoou& 71 6 Activity patters of Camponotus acvapimensis 72 6.1 Introduction • • • • 72 6.2 Materials and methods • • * • 75 6.2.1 Foraging activity .. * • • » 15 6.2.1.1 •I Diurnal activity pattern .♦ •• I • 0 75 6.2.1 .2 I Nocturnal activity pattern *• • • 76 6.2.2 Foraging space and distance • • 77 6.2.3 Induced foraging •• •• • • 73 6.2.4 Ants and phytophagous insects associationship 75 6.2.5 Flight activity . • • ♦ • • 7? 6.2.6 Food items collected by ants •• • • 7? 6.3 Results/ Observations • • SO 6.3.1 Foraging activity of Camp •acvapimensis 30 6.3.1.1 Diurnal activity . • • • 81 6.3.1.2 Nocturnal activity • * 04 6.3.2 Induced activity .♦ * • 86 6.3.3 Foraging space and distance •• • o 86 6.3.4 Ants - phytophagous insects associationship 88 6.3.5 Food items collected by ants •• O 0 83 6.3.6 Flight activity • • ♦ • o » 8Q University of Ghana http://ugspace.ug.edu.gh - X p a g e 6.4 Discussion .. .. .. 9l 6.4.1 Foraging activity of Camp, acvapimensis 9i 6.4.1.1 Diurnal activity ., ,, 91 6.4.1.2 Nocturnal activity Jt #> 94 6.4.2 Foraging space and distance 95 6.4.5 Food items collected by Camp, acvapimensis 97 6.4.4 Ants and phytophagous insects associationship 99 6.4.5 Flight activity 100 7 SummaryJ »• •• •• 102 8 References s--• • »• •• Appendix Tables. 107 University of Ghana http://ugspace.ug.edu.gh - xi - a b s t r a c t The distribution, nesting habits and activity as well as seasonal population pattern of Camponotus acvapimensis Mayr were studied in a savanna area of Ghana* The ants were disti- buted in the poor shaded areas of the savanna grassland* Type of soil, light and food availability were found to influence the nesting and antis distribution. Analysis for preference of nesting sites indicated that the ants nest in the soil or grass roots but where wood is available, the bark formed a very good nest. Two types of nests were identified. The first type is the primary nest with many cells. Such nests contained the highest populations and had larvae, pupae, workers, soldiers and alates in them. The second type is the secondary nest that largely sejyed as nests for tending other insects and such nests had few larvae and pupae and rarely alates with the soldiers and workers tending the sap feeders. The ant is active both day and night but peaks of activity were comparatively high during the nights and lew during the day. The workers were numerous and formed a maximum of JOy0 or more of the ants (excluding larvae and pupae). The workers and soldiers were found to forage all day but the number of soldiers foraging at night was comparatively few. The ants feed on sugary materials such as nectar and honey dew from other insects. They also collect flower parts and vegetable tissue. The ant tends aphids, coccids, other homoptera and hemiptera. University of Ghana http://ugspace.ug.edu.gh C H A P T E R O N E GENERAL INTRODUCTION University of Ghana http://ugspace.ug.edu.gh C H A P T E R 1 GENERAL INTRODUCTION West Africa possesses a rich fauna of ants that are found in different ecosystems* The distribution of these ants tends to be influenced by factors which are mainly environmental. Vegetation cover plays an important role in habitat selection of ants as plants provide the source of food and nesting sites for most of them. Two types of vegetational zones are recognizable in Ghana. These are the forests which are made up of rain forest orIcocoa forest! (Leston and Hughes 1 968) and the savanna grassland. In Ghana many genera and numerous species of ants are found. Room(1971) found 48 genera and 128 species of ants on mistletoe which made him conclude that about 50% of the insedt species found on cocoa mistletoe Tapinauthus bangwensis (Ehgle and )C Krause) were ants. TJxe occurrence and activities of ants have thus been studied by several workers (e.g. Leston, 1968, 1 971 Aryeetey, 1971J Majer, 1972, 1973, 1974, 1 976a-c,# Ackonor, 1977? Adabie, 1977)# These works concerned ants in the cocoa ecosystem only. No work appears to have been done on savanna grassland ants which have become economically important due to the introduction of large scale farming in the grassland areas of Ghana. The distribution of the savanna species has only been mentioned in the work of Majer(l972) where he placed - 1 University of Ghana http://ugspace.ug.edu.gh - 2 Camponotus acvapimensis Mayr under the sub-dominant group Krhich under certain field conditions may numerically predominate and exclude all forms and species! • Room (1 971) mentions the nesting sites and materials used for nesting by Camp* acvapimensis. Goetsch(l 929) records the seed collecting habits of some ants and this is an indication that Camp, acvapimensis may have preference for the new food source which may be grown as a monoculture* The present work is an attempt to study Camp* acvapimensis in the savanna ecosystem* Although the species Camp, acvapi- memsis has Ghana as its type locality, no work has been done ©n the species. The only findings that have been made on the species are the incidental observations during investigations on pests of cocoa or other ants, or collecting ants and insect species in the cocoa farm ecosystem* These include the work of Gorenz (1969) who reported that Camp* acvapimensis was the possible cause of the pod-rot disease carrying the inoculum from the base of the tree to the pods in the canopy. Evans (1 971, 1973) confirmed the findings of Gorenz and .accepted the role of the ants in the transmission of the spores of the fungus from the soil to the pods in the canopy* Evans(1 971 , 1973) felt that the spores might have been carried on the bodies of the ants* Apart from these reports on the involvements of the ants in fungus transmission, some reports have also been made on the foraging and nesting habits. Bigger(1972) collected the ants University of Ghana http://ugspace.ug.edu.gh - 3 - on cocoa trees,’ Room(1 971) recorded the presence of the ants,, nesting and foraging areas in the cocoa farm as well as collecting the ant from Tapinanthus bangwensis. Firempong 0975) reported on this antis association with the aphid Toxoptera aurantii(Boy.) and other scale insects, and Majer (1972) mentioned the ant as preferring broken canopy in the cocoa ecosystem. Mayr(l865) described the characters of the workers of Camp, acvapimensis but no account is available of the castes of this species* Whe*ler (1922) however, mentions that the species of Camponotus often form populous colonies and exhibit great diversity of nesting habits. He also states that they may live in the ground either under stones or in crater nests and others under bark in dead wood and hollow twigs. On Camp, acvapimensis, Room(l97l) recorded the antis nesting habits and indicated that the nests are found within the soil or in dead wood in the soil. In all these, no details of the worker castes are available. Wheeler(1922) mentioned that the species from Bolenzi in the Republic of Congo were found nesting in the trunks of Oil Palm trees. Very little is known about the behaviour of the species but Majer(l 972) mentioned that it may exclude all forms from areas in the cocoa farm edosystem without shade. This could imply numerical abundance and pugnacious habits. Very little is known ^bout the food habits of the ants but several workers such as Firempong(l 975) , Evans U 971, 1975) and others have ±ound tha,t the ant tends aphids, scale University of Ghana http://ugspace.ug.edu.gh insects and mealybugs in the cocoa ecosystem. Wheeler(1 922) also reported capturing some workers of Gamp, acvapimensis at Faradje in the Republic of Congo while tending plant lice on the young leaves of orange trees* Evans(1 97^ > 1973) mentioned that these ants used gnawed materials for building tents ovsr the mealybugs which they tend. Predatory activity of the ants has not been reported by any worker but Wheeler(1 922) found the workers of Camp, acvapimens i s in the stomachs of the toads Bufo regular is Reuss and B. fenereus Boca^ ge. One major worker was found in the stomach of a frog, Rana occipitalis Gunther, showing toads and frogs prey on these ants* The aim of this is to provide information on the ecology of Camponotus acvapimensis* Distribution of the ant in relation to farming habits as well as the effects of external factors such as vegetation are studied. The type of nests constructed by the ants, their colony size as well as the phenology of the caste and brood over the period of study are examined. Activity pattern, foraging range and distance and the flight activity are considered. The antis association with other insects is investigated. Attempts are also made to furnish information on interaction with other ants occuring in the area. University of Ghana http://ugspace.ug.edu.gh C H A P T E R T ¥ 0 G E N U S C A M P O N O T U S M A T E University of Ghana http://ugspace.ug.edu.gh COTUS flAJviPONOTUS MAYR The genus Camponotus was erected by Mayr(l862) and the number of species in the genus has increased with time. Smith (1866) noted the numerous additions and remarked 11 the number had increased from 18 in 1861 to 32 in 1865"- He suggested that new species were likely to be discovered# The discovery of new species did indeed take place and numerous new species vrere described by Emery(l883)j De Geer(l884); 0ertzen(l887) and Forel(l890)• The sub-divisions of the genus towards the end of the last century was caused by the large number of known species# Emery(l896) sub-divided Camponotus into more or less natural groups of 26 subgenera but Forel(l912) sub-divided the genus into 20 subgenera and in 1914 published a list of all known species at that time# Emery(l920) also published a revised classification of the genus taking into account geographical distribution of the species. The former status of Camponotus as belonging to the subgenus Hyrmoturba by Forel(l9l2) was followed by Emery(l920}# Wheeler (1922) described the genus Camponotus as a "huge cosmopolitan genus, comprising more than 1,000 described forms'1. He noted the frequent occurrence of the species of the genus Camponotus in all countries excepting United Kingdom and New Zealand# Wheeler(1922) further attributed the University of Ghana http://ugspace.ug.edu.gh extraordinary variability of many of the species of Camponofag. to their response to slight differences in their environment. He. recognised too different foms of CajnEonotus grouped by their appearance and. habits. The first group was described as smaller forms that are stolid, apathetic or timid( C amp on o tus acvapimensis belongs to this group as it is very reluctant to bite, although possessing quite formidable mandibles). The second group comprises maxima workers of the large species belonging to the subgenera Dlafflflyrmex. Hyrmoturba, Hyrmothrix and Myopiromis which are pugnacious and are capable of ^inflicting wounds with their powerful mandibles. The subgenera of Qamponotus described by Wheeler (1922) are distributed within certain areas of Africa and A£ia as follows; REGION STJBGENERA Ethiopian Kyrmoimrba. Dinomyrmex. Myrmosericus. Hyrmthrix. (frthonotomvrmex. Mymotrema Mvrmopyromis. Evrmorhachis. Nyrmops amirm. M.yrmamblys and Colobosis. Camponotus. Myrmosaulus, Mayria. Malagassy Hvrmonesites and M.vrmopytia . University of Ghana http://ugspace.ug.edu.gh - 7- REGION ~ “ SUBGBNERA African jyryrmonsamma, Kvrraopiromis. Madagascar Mvrmosa^a. Ma.vria* llvrmonesites and Mvrmopytia. Mvrmoturba. J^yrmosericus. and West African Orthonot omyrmex. Southern China across India to Myrmosericus. and Myrmoturba. South .Africa and Gulf of Guinea Table 1# Distribution of subgenera of Canrponotus (after W&eeler 1922) Out of these species (Table 1), Wheeler(l922) named a few species that have a greater coverage with respect to distribution. These are Cannponotus (Myrmopytia) species that occur in the MJ£Lagassy region* He names Cam-ponotus (Myrmoturba) maculatus (Fabricius) as the typical form which is West African and two other species C^aponotusfayrmosericus )rufo>?lacus Jerdon and Camponotus(Orthonotomyrmex)sericeus (Fabricius) as having singular distribution with forms of Camp, maculatus occxiring in all the continents* Camp, rufOfflaucus lias many varieties that are found in Southern China across India and Equatorial areas to South Africa and Gulf of Guinea. University of Ghana http://ugspace.ug.edu.gh Wheeler(l922) mentions Camp* sericeus as occupying similar ranges but showing little tendency to produce sub-species and varieties# Later works on the species included changes in spellings, for example, Camp* acwapimensis; Camp* a&qwapimensis; Camp* akvapimensis and Camp* akwapimensis Emery(l877; 1899)j C 905)* The type locality of the species is from the Akwapim Mountains of Ghana and this might have caused confusion in the spellings of the type species. In Ghana, the species occurs in several places and these include Aburi, Tafo-Akim, Bunsu, Kadef Mampong Akwapim, Kumasi and other places (Evans, 1971; Room, 1971; Firempong, 1975; Majer 1972, 1973, 1976a - c). These places and thd type locality of the species are shown on the map (fig l). In Ghana, three species of the ground nesting ants of the genus Camponotus are common in grassy patches in the saxanna areas or clearings in the forest zones but as many as 10 species of Camponotus were recorded by Room(l97iJ during a sampling programme in a cocoa farm. The three grassland species found in Ghana and nesting in the soil are acvapimensis Mayr, Camp, chrysurus Gerstaecker and Cag£* vestitus (F.Smith)* In terms of numbers, Camp. _a_cyapimensis is by far the dominant species in the grassy patches followed by Camp, chrysurus and Camp, vestitus. the last preferring more or less a bare ground* - 8 - University of Ghana http://ugspace.ug.edu.gh FIG: 1: Map of Ghana showing distribution of C. acvapimensi s I ^ Akwapim Hills @ Togo Ranges University of Ghana http://ugspace.ug.edu.gh C H A P T E R T H R E E rpTTE 3 TODY APJSA 3.1 LOCALITY The study area was situated around Presbyterian College and Madina (fig 2). The different plots (A and b) were selected for the study* The plots measured about 2 hectares and "ere continuous. They were boardered (fig. 2) on the far !?orth by the Madina new road, on the far East by the Institute of Professional Studies buildings (l.P#Sf), on the immediate South by the road to I.P.S. and on the immediate West by the bungalows of Presbyterian College (fig* 2). Within the idiots the vegetation covers consisted of : (i) Plot A - an area frequently disturbed by fires when the grassland is dry during certain periods of the year. It was largely de^ ip'd of green vegetation during- the period when conditions were not favourable and with green vegetation when conditions were favourable. (ii) Plot B — an area not frequently disturbed and had cultivated plants such as. com, cassava, and cocoyam, and was not subject to fires. 3^2 Plot A. (savanna grassland) This was a fire frequent plot because during the dry season® (late December to March or April and late July to early September), the grasses dxy-up and the bush fires start sweeping through the field, consuming the entire vegetation. The fires are set "by the farmers: as frequently as the grasses dry-up and once started* they University of Ghana http://ugspace.ug.edu.gh KEY TO FIGS: 2, 4 & 5 CONTINUOUS SHADE BROKEN SHADE © POOR SHADE 0 NO SHADE © t r e e © NEST OF £. S ER IC E IJS T Z ^ n e s t - OF C. 0 NEST OF C. ACVAPIMENSIS c f .. „ „ (frj n <• <■ A " “ DRIVER ANTS -G- PLATYTHYREUS 1 cm. = metres IN SOIL " STUMPS *• TREE University of Ghana http://ugspace.ug.edu.gh ftocx VAo.9 s\\0* \r\Q s'o^e a u v *\oo ot o n ,lot O0 University of Ghana http://ugspace.ug.edu.gh - 1 2 - The shrubs are reduced to mere sterns and beac very few (if any) leaves. Only trees and few shrubs have green leaves (table 1). 3*5 Plot B (farm-1 and) The meteorological factors affecting this plot were the same as those affecting Plot A. However, there were major differences caused by the different shade regimes provided by the plants such as Zea mays L. (maize), Manihot esculent a Crantz (cassava) • Musa paradisiaca L. (plantain) , with the last forming the dominant plants of the area. The seasons influencing Plot B were the same as those influencing Plot A. The main differences were caused by the shade provided by the shade plants in the area as well as the ability of the cultivated plants to survive periods of drought. The grasses are absent or when present they are found in pathhes in few places whilst in their places are different plants, Humidi'ty ranges from an average minimum of about 5&/q R.H, for the periods of day to about 98/q - 9 % R.H, during the rainy periods. Sunshine varies with the seasons but the effects of light intensity on the soil depend to a greater extent on the type of plants growing in the area as dominant species and those growing as an undercover. The shade depends on the type of plants found in the area and in particular where plantains and cassava are grown, the shade is quite . .,:&ense, Tress are absent which is the result of removal of such plants during the clearing University of Ghana http://ugspace.ug.edu.gh University of Ghana http://ugspace.ug.edu.gh Ta bl e 1 • Di st ri bu ti on of pl an ts on Pl ot A, w0 •H £5 oo •tjS cdro N S -=£j w cS b -p Q •H rO CtiP. cti o H X rH cd 8 G ctf <1) O rC 03 •H •rH rd g & co cti c I—1 •H g Pi (D 1 o* INTRODUCTION Various workers have studied the ecological distribution of ants in different habitats in several ant species ( for example, Gosswald, 1932; Tabolt, 1934; Dennis, 1958; Leston, 1971; Majer, 1972, 1973, 1974, 1976a-c). In all these cases, selection of habitats was discussed as being influenced by climatic factors and conditions such as the availability of food and vegetation cover, Camponotus species have been reported from several eco­ systems# Brown(l959) reports Camponotus species associated with other ants on coconut plantations on Solomon Island* He mentions four different species of ants but specifically he stated of Camponotus species as foraging on palm stems but not on their spadices. Wheeler(1922) records Camp# acvapimensis from the following places where the ants have been found: GHANA : Akwapim Mountains, GUINEA: Eindia; Camayenne; Mamou; Los Island. SIERRA LEONE: Most places* LIBERIA: Grand Bassa ; Junk River (H# Brauns), SOUTUKRN . .NIGERIA, ; Moor plantation near Ibadan; Old Calabar. CAMEROON : Victoria. .Ti^ J^jAjSA : Banza Masola; Ganda Sundi; Mondombe; Congo da Lemba; Tambui; Zambi; Faradje; Garamba; Bolengi; near coquilhatville; Stanleyville; Vankerckhonville; University of Ghana http://ugspace.ug.edu.gh - 1 6 - Akangej Niangara; Thysville and Bala. RHODESIA s Bulawayo. KENYA AMD TANZANIA : Kigerama; Mombasa; Tangan; Mto-ya-Kifara; Kilimanjaro and Patta Manda. UGANDA : Chao ans engula • SOMALIA; Lower Gunaya; Giar Buie; between Obbia and Berbera* ETHIOPIA : Arussi Galla; Ganale Gudda; Dime to Bass Narok; Bela; Coromma; Western Ethiopia; Keren and Asmara. In Ghana the species hatfe been collected from several areas and these include places such as Tafo, Bunso, Eumasi, Akwapim Mountains (Mampong) and Kade. Hajer(l972) in a series of samples collected Camp. acvapimensis on cocoa, trees at a frequency of 31 out of 144 samples* Their occurrence and presence in different places and localities have also been reported by Room(i974.) who collected Camp. acvapimensis on a mistletoe Tapinanthns bangwensis In another work, Room(1971) names Gamp* acvapimensis as one of the ants found under cocoa farms and foragiiig on herbs under the cocoa trees. “ivheeler(1922) described the genus as a large group with mo^e than 1f 000 described forms, and one is tempted to investigate the status of the species of this genus within their natural ecosystem, as to whether their relative abundance will make them dominant or not. With respect to numbers, Leston(i971) recog­ nized the ant species of the cocoa farm to belong to 3 distinct groups whieh lie named as 1 (1) Dominants (2) Co-dominants and (3) Hon-dominants. Majer(l972) considered £ajnp. acvapimensis as University of Ghana http://ugspace.ug.edu.gh a sub-dominant group and further stated that ”under certain field conditions they may numerically predominate to exclude all other species from the area of cocoa farm ecosystem which are devoid of shade and having characteristics of savanna grassland". Wheeler(l922) however, refers to the genus as Moften forming populous colonies and exhibiting great diversity of nesting habits’1. This is an indication that any of the species within a favourable ecosystem can reach the status of1 dominant1 species. The status accorded to Camp* acvapimensis as a sub- dominant group by Major(1972) applies to the species only in the cocoa farm ecosystem which is possibly not their natural habitat. Wheeler(l922) emphasizes that the "extr^rdinaiy variability of many of the species in response to slight differences of environment makes the genus one of considerable interest to the student of geographical distx*ibutionfi. Several reports on Camp, acvapimensis in the cocoa farm eco­ system indicate the importance of these ants within the ecosystem. The aim of this section is to map the nests of Camp, .acvapimensis in relation to shade regimes as well as in relation to the other ants nesting in the area* ITesting habits of the ants will be presented and the percentages of such nests in the habitat determined* The colony size of the ants will be investigated and the number of nests forming one colony would also be worked out. - 17 - University of Ghana http://ugspace.ug.edu.gh 4.2 METHODS 4.2.1 of nsstg and aest distribution in relation to shade, regimes on Plot A. (Savanna) AA area mostly covered with grasses (referred in chapter 3 as stud# plot A) was demarcated. Trees and other plants in this area were recorded and those bearing nests of Cam_£*acvapi- memsis were mapped out* Distribution of nests was studied and consideration given to the place where the nest was found, preference of nesting area was determined by the abundance of nests in a particular area. A comparison table was constructed to determine if there was any significant difference in the distri­ bution of nests on the savanna area as compared to the farming area* The entrances to the nests were mostly found by observing and tracking the ants that were carrying food materials. Another method used was to follow any ant that had its abdomen (gaster) raised and moving very fast due to excitement into its nest and then labelling the nest. 4*2o2 Abundance of nests and nest distribution in relation to shade regimes on Plot B(farm land) A search for nests of Camp, acvapimassis was done in the farming area(plot b) which had plantain, cassava and corn during the wet season and no corn during the dry season. Counting of the nests was done by observation and searching through the plant­ ain field. The plantains with nests were noted and their- numbers tabulated. Nests found in the soil were distinguished - 104- University of Ghana http://ugspace.ug.edu.gh from those found on the plantain suckers by using different symbols. The effects of the changing seasons on the abundance of nests were tabulated and the percentage distribution of such nests also determined. The percentage distribution of such nests with respect to shade regimes was calculated for the plot during the wet and dry seasons and the results tabulated© Other ants nesting in the area were mapped and the percentages of their nests in different shade regimes was obtained. This was then compared with the nestr . distribution of Cam p. acvapimens is in relation to shade preferences. Investigation into shade preferences was carried out by three methods. (1) By counting the number of nests in the different shade regimes and finding out the percentage of such distribu­ tion in relation to the other shade regimes; (2) By artificially shading areas having numerous nests and determining the effects of the shade on the nesting preferences of Camp, acvapimensis: (3) By clearing the shade provided in some areas and finding out if the nests present will be abandoned or other nests will be established in such areas. The effects found were recorded and preferences estimated by abandonment or establishment of nests in such areas. Similar calculations were made for the other ants with reference to the number of nests found in the soil and those found on plantain suckers or other places. ~ 19 - University of Ghana http://ugspace.ug.edu.gh -20- 4.2.3 Colony size Colony size was determined by the following methods: 1. Transfer of workers foraging on leaves Camponotus flcva-pimensis is timid biit easily gets excited and moves away when disturbed. As a result, the ants could not be induced for transferrence to other places by 8ny easy method. The only way to cause an easy transf errence of the ants to other areas was to carry them from where they were found foraging. The transferrence was done gently to avoid excitement. By means of a sharp knife, the branch on which the ants were foraging was cut and carried carefully to an area with other ants of the same species foraging and gently putting the cut branch on the vegetation and observing the interaction of the ants in the new habitat as well as the ants th&bwere found there. The behaviour of the ants was then recorded. 2. Attempted excavation of some nests A few centimetres of some nests were excavated and the ants disturbed. The ants were observed to find out if they would abandon their nests to establish new nests. The behaviour of the ants such as the carrying of larvae and pupae was then observed and recorded. The new nests where the larvae and pupae were sent were considered as parts of the colony. 3. Prftvidinff a bait Since Camp. acvapimensis workers tend aphids and other Homoptera for honey dew and other sugary materialsj Sugar University of Ghana http://ugspace.ug.edu.gh baits were used as a source of enticement. The ants in the area were then observed during their feeding habits and their reac­ tions towards each other recorded. In this method it was assumed that' these ants being timid may avoid ants of the other colonies and would move off rapidly. It was also assumed that the ants from the same colony may feed with the least disturb­ ance from other ants, from different colonies. Characteristics such as moving towards the nests are characteristics that were used to determine acceptance and rejection of a member of a colony by another member. 4*2.4 Distribution of Camp, acvapimensis in relation to other ants on plot A. The mapping of the nests followed the same method as the one used on plot B. The only difference was in the mapping of the nests occurring on the plantain suckers. Other nests in the same places as those found on plot B are shown with the same symbols as in plot B in order to bring out the similar- lit. as of nesting habits. Ants nesting in the soil and grass roots or shade trees were also mapped out. The occurrence of such ants was subjected to statistical analysis to see if there was any significant relationship between the occurrence of these ants and Can P. acvapimens is. The presence of such nests was plotted on the same map with different symbols in order to differentiate them from those of Camp, acvapimensis. In constructing the maps, the area of the plot was - 21 - University of Ghana http://ugspace.ug.edu.gh measured and the trees and other features that were found were plotted using the same scale. Distribution of Camp, acvapimensis inafc relation to other ants on plot B. The other ants nesting in the area Plot B were mapped out and the distance of their nests to those of Gamp, acvapimensis also determined. The number of dominant ants was recorded and their nesting habits discussed in relation to the nesting habits of Camp, acvapimensis. A search for the ants was carried out by -methods similar to those used for Gamp, acvapimensis. The ants in some cases were provided with crumbs of bread which they carried into their nests. The ants were then identified and numbered together with those of the same species. Their nests were also numbered. By this method, almost all the ants that were dominant species in the area were known and their associationship with Camp* acvapimensis found out by considering their interactions with Camp, acvapimensis. The nesting preferen­ ces of all the ants that were found were considered. Those ants that were found on plantain suckers with and those without nests distinguished. The common ants in the field were collected and identified to genera with Boltonls (1973) key. OBSERVATIONS / RESULTS Abundance of nests on Plot A The number of shade trees in the area are shown in fig 4* (i) wet season During the wet season, the nests were found at 3 - 22 - University of Ghana http://ugspace.ug.edu.gh d&fferent places — Nests in the soil:- These nests were found as entrances to the main cells underground* Nests in frrass stumps and roots;- These nests were found in the roots of growing grasses and others occurred between the stalks of grasses and others In the middle of foliage leaves (Pig*6)« Nests in dead wood:*- These nests were in dead wood or dead portions of living trees# They had one or more entrances# Out of a total of 54 nests found during the wet season, only $ were in trees* A total of 15 (27*850 were found in the roots of grasses arid the remainder 36 nests (86?£) were in the soil# Of a total of 12 trees found in this area, only 3 had the nests of Gamp#acvapimensis# This number represents 25^ of the nest bearing trees# Efcom the above observations it seems evident that these ants have preference in nesting in the soil as compared with nesting in wood (table 3)« (ii) Ifery season The results obtained on nest distribution during the dry season on plot A differed slightly from the results obtained in the wet season# Of the 12 trees in the area, only 3> (the same trees that were found during the wet season) had nests of Gamp♦ acvapimensis « This figure represented 255^ nest bearing trees# The three types of nests were distributed as follows: - 23 - University of Ghana http://ugspace.ug.edu.gh Table 3: Distribution of nests of Camp.ac vap imens is during the wet season on plot A (a) Number of nests on trees and percentage nest trees Total No. of trees Trees with­ out Nests Trees with Nests. % trees with nests Nest on trees 12 9 3 25 (b) Percentage distribution of nests in different nesting area Locality Total number of nests % Nest distribution In soil 36 86*0 In grass, 15 27.8 On trees 3 5.6 Total 54- 100.0C?' (c) Percentage distribution of nests in various habitats Habitat Total in trees in passes in the soil (shade _ regime) nests No. No. % No. cl No shade 4 0 0 3 75 1 25 Broken _sh§de_ 17 0 0 8 47 9 52.9 Poor shade 23 0 0 1 4.4: 22 95.7 Continuous shade 10 3 30 3 30 4 40 University of Ghana http://ugspace.ug.edu.gh 3 (d) Percentage distribution of soil nests in diffea:ent shade regimes on plot A in the wet season* Habitat Ho* etf nests % distribution No shade 1 2.8 Broken shade 9 25*0 Poor shade 22 61.1 Continuous shade 4 11.1 Total 36 100*0 (e) Percentage distribution of nests in grass roots in different shade regimes on plot A in the ?;et season. Habitat Ho. of nests % distribution No shade 3: 20.0 Broken shade 8 53*3 Poor sha.de 1 6.7 Continuous shade 3 20*0 Total 15 100*0 (f) Percentage distribution of nests in trees in the different shade regimes on plot A in the iret season. Habitat Ho* of nests % distribution No . shade 0 0 Broken shade 0 0 Poor shade 0 0 Continuo us shade 3 100*0 Total 3 100.0 University of Ghana http://ugspace.ug.edu.gh Table 4: Distribution of nests during the wet season on plot B. (a) Distribution of nests in different nesting areas Locality total no of nests $0 distribution In soil 38 53.52 nests on suckers 33 46.5 Total 71 100.0 ( 0 Distribution of nests ±n the various habitats in the %et season Habitat No.of,suckers No .with nests No. without nests % of suckers with nests No shade J 1 2 33.3 Broken shade 29 19 10 5 7 .6 Poor shade 6 3 3 9.1 Continuous sha.de 100 10 90 30 .3 Total 138 33 105 100.0 University of Ghana http://ugspace.ug.edu.gh 4 (o) Percentage distribution of nests in various habitats and localities Habitats Total nests in -plantains in ^rass roots in the soil No* e//W No* ti No. q;jQ Bo Shade 3 1 33.31 0 0 2 66.7 Broken shade 21 19 90.5 0 0 2 9*5 Poor shade 31 3 9.7 0 0 28 90.3 Continuous shads 16 10 62.5 0 0 6 37*5 (d) Percentage distribution of nests in the different shade regimes on plot B in the wet season Habitat No. of nest 5o distribution No. shade 3 4.2 Broken shade 21 29.6 Poor shade 31 43.7 Continuous shade 16 22.5 Total 71 100.0 University of Ghana http://ugspace.ug.edu.gh Table 5ai Comparison of % distribution of nests of Carrrp»acva’pimensis in the soil and other places on plots A and B. Nest in the soil Nest in other area Habitat Plot A % PlotB a Plot A o'A Plot E 5* No shade 1 2.8 2 5.7 3 16.7 1 3 Broken shade 9 25 2 5.3 8 44.4 19 57.6 Poor shade 22 6 m 28 73.7 1 5.6 3 9.1 Continuous shace 4 11.1 6 16.0 6 33.3 10 3a3 5 b. Comparison of n st distribution during v«fet and dry seasons on Plot A. Locality DRY SEASON ''Tr-'in SELiSOI! No. oC/v No. e' In trees 3 9.7 3 5.6 In soils 26 83.9 35 86.0 Grass roots 2 6.5 15 27.8 Total 31 54 University of Ghana http://ugspace.ug.edu.gh T&ble 6: Percentage distribution of nests on plot A in the dry season: 6 (a) Nest distribution on trees No* of nests No. of trees trees with c/j nest without nests nests trees. Nesting on 25trees 12 9 3 6 (b) Percentage distribution of nests in various localities on Plot A Locality total no. of nests $3 Nest distribution In soil 26 83.9 In grass root 2 6.5 In trees 3 9*7 Total 31 100.0 6 (c) Percentage distribution of nests in various shade regimes HABITAT (shade regime) TOTAL IN TRESS BT GRASS ROOTS U! SOIL Nests No. o< No. 0' No. o' No shade 30 0 0 2 6.7 28 93 .3 Broken shade 0 0 0 0 0 0 0 Poor sh^de 0 0 0 0 0 '0 0 Continuous shade 4 3 5.0 0 0 1 25.0 University of Ghana http://ugspace.ug.edu.gh Table 7 s Percentage distribution of nests in various habitats on plot B during the dry season: 7 (a) Percentage distribution of the nests on plantain Habitats No. of suckers No* with nests No* without nests % with nests. No shade 3 1: 2 Broken shade 29 16 13 Poor shade 6 3 3 Cont* shade 100 5 95 Total t 1 — * 1 Lo 03 1 1 1 1 1 f 25 rov—r— II 7 (b) Percentage of nest distribution in various localities LOCALITY Torn JS mSS DISTRIBUTION In the soil 22 45.8 on suckers 26 54*2 Total 43 100*0 University of Ghana http://ugspace.ug.edu.gh 7 (c) Percentage distribution of nests with respect to shade regimes HiUBITAT Total on suckers in grass in soil No. % No, 0/A No. c'/•-* No shade 3 1 33.3 0 0 2 56.7 Broken shade 19 16 84*3 0 0 3. 15.8 Poor shade 17 3 17.7 0 0 14 82 .4 Cont. shade 9 6 66* 7 0 0 3 33.3 7 (d) Percentage distribution of nests in the differ ent shade regimes on plot B in the dry season Habitat Ho. of nests % nest distrxbutic No. shade 3 62.5 Broken shade 1? 39.6 Poor shade 17 39.4 Cont, shade 5 18.8 Total 48 100.0 University of Ghana http://ugspace.ug.edu.gh 8 (a) Percentage distribution with respect to shade regimes Table 8: Comparison of the percentage distribution of nests on the two plots A and B. on plot Jb Habitat Wet season dry season No. % Ho. % No shade 1 2.8 28 96.6 Broken shade 9 25.0 0 0 Poor shade 22 61.1 0 0 Continuous shade 4 11.1 1 3.5 Total 36 100.0 29 100.0 8 (b) Percentage distribution of nests with respect to shade regimes on plot B Habitat Wet season dry season No. m the observations, nest distribution seems to be related to shade regime* Most nests were found in the soil but not in dead wood in the soil* This contradicts Room (1971) who reported that the ants nest in dead wood in the soil* Of the total number of 48 nests that were dug during the study period, none were in dead wood in the soil* Nesting in the wood was found to be only a matter of availability. It is therefore not surprising that Room (1971) found the ants nesting in dead wood in the forest areas where there was plenty of wood in which the ants could nest* In this, it is also expected that some nests would be in dead wood in the soil* With respect to shadS regime, it was found that the ants were more in the poor shade regime area* The number of nests occurring in the area constituted $1 *1^ of the total - 34 - University of Ghana http://ugspace.ug.edu.gh nests and this tends to agree with the findings of Majer (1972) that Gamp. acvapimens is under certain conditions becomes dominant in areas of forests that resemble the savanna ecosystem. The savanna ecosystem that he referred is typically the poor shade regime of this study. The presence of nests in the continuous shade regimes was only found to be due to the preference that the ants had for nesting in wood as compared to nesting in the soil# Distribution of nests in grass roots had more of the nest3 in the broken shade regime. This is likely to be due to the fact that the poor shade area had most of the grasses reduced to mere roots when the light intensity becomes very strong on the roots and causes the ants to abandon such nests* The nests are therefore established in the broken shade regimes to prevent the sun from drying up the eggs* In general, the ants do not nest in shadeless areas and they do not prefer continuous shade areas either* Preference for shade regimes therefore was for poor shade areas follov/ed by a broken shade area. In the dry season however, there were more nests in the shadeless area. This is because the grasses are often burnt and the nests found at the grass roots as well as in the soil in the grass covered areas become bare and the area is therefore mostly a ‘no-shade*area* University of Ghana http://ugspace.ug.edu.gh - 36- 4«4*2 Abundance of nests on Plot_B On plot B the findings were different. Distribution of nests in the wet season followed a different pattern in that more nests were found in the poor shade area but the continuous and broken shade areas had almost the same percentage distribution. Camp.acvapiaensis thus nests in the poor shade areas but when farming practices permit the area to be changed to continuous or broken shade areas then the ants will be more in the broken shade areas and a few more in the continuous shade regimes. In the dry seasons the ants are found in the same broken shade areas and they are found to move more into the farm lands since the shade in the savanna areas is reduced considerably. 4*4»3 Distribution of the nests Nests of the species occur mostly in the soil* G&mp.acvapimensis is therefore a ground nesting ant with little tendency to nest in dead wood. They also construct secondary nests to tend aphids, scale insects, some Homoptera and Ooccids on some vegetable crops such as plantain and melons. 4*4*4 Colony size The ants were not found to form large colonies and the distances between such colonies increased as the animals constructed nests far away from their primary nests in order to tend other insects. University of Ghana http://ugspace.ug.edu.gh - 37 - Cblonies may be separated by longer distances or the colonies may be very near the primary nests depending on the distance between the place where the plant on which they mat to tend the insects is found. Members of the same colony may feed together without much excitement excepting when they come in contact with members of the other colonies for the first time. T&ien they belong to the same colony, they forage or feed from the same source of food but some of them withdraw to other sides for foraging. Ants of the same colony do not move away from the food source into their nests to recruit more foragers or soldiers when they come in contact with members of the same colony. A. difference exists between ants of one colony and those of different colonies. Those ants belonging to different colonies show antagonism which they indicate by quick dashing with open mandibles at ants of the other colonies. Some of the ants move very fast, away with raised gasters in undetermined directions and many such ants entering into their nests and later coming out of the nests with other workers or soldiers castes to the source of food. Gamp, acvapimensis is liMy to produce pheromone for warding off the ants from a different colony. As many as 7 nests, both primary and secondary were fotmd to Constitute one colony. Colony size of the speckles requires further investigation as the ants are timid and are very difficult to control in University of Ghana http://ugspace.ug.edu.gh - 38 - experiments that were conducted for the determination of colony sizes. Further work on this ant should consider the primary and secondary nests as well so as to he sure that such nests are part of a primary nest. 4.4 .5 Distribution in relation to other ants The nests of the other ants found in the study areas were very few and this did not make the other ants dominant in terms of presence of nests in the nesting areas. However, in terms of numbers Camp, sericeus were found to he more populous than Camp• acvapimensis. The numerical consideration of foraging workers for a given period of the ants indicate that Camp, sericeus are more a.etive during the day than Camp»acvapimensis. The differences could he due to the fact that Camp, sericeus forages only during the day whilst Camp»acvapimensis forages during the night and day constantly with little breaks when the light intensity becomes high. These two ants were found to have preference for the same nesting areas but were not found to be involved in fights but Camp.acvapimensis was found to attack Camp. serioeus wherever the two came in contact with the latter retreating from the area# Both ants are timid as compared to other ants described as aggressive. Ackonor (l977) "by a series of experiments established that Cataulaucus ffuineensis were very aggressive and involved even members of other colonies in fights resulting in loss of parts of their limbs.' University of Ghana http://ugspace.ug.edu.gh 4*4*5 * f Gamp.acvapimensis and Camp.sericeus Nests of these two ants were found in the same areas* TThere trees were found on the plot, Canp • ac vap imen sis was found to nest in the dead portions of such trees but the nests were more confined to the soil in such areas# Canrp * ac vap imen sis is therefore a ground nesting ant and was not competing with Caanp.seiceus for nesting sites in the wood. The 3iests of Gamp »acvap imen sis were found in the bark of the tree but Camp.sericeus made holes into the wood as nests. With respect to shade regimes, the two ants were found to have preference for areas with broken and poor shade. As the two species have different nesting sites and do not fight but only Gamp.acvapimensis shows a tendency of attacking at times, competition for nesting sites was not noticed but the ants were found to compete for food source as they all show preference to sugar diet. Co-existence is therefore more mutualistic thaji antagonistic. These were the two dominant ants occurring in the savanna region under investigation. 4.4* 5.1.1’Numerical abundance In terms of numbers, the workers of Carnp+sericeus were found to be more at any given time as compared with Camp. acvapimensis. The number of ants found during a period of 10 minutes coming out and entering the nests of the two ants (table 12) gives an indication that Camp.sericeus is numeri­ cally abundant as compared with foragers of Gamp.acvapimensis, - 3 9 - University of Ghana http://ugspace.ug.edu.gh 4*4#5* 4q4<>5* This difference might he due to factors other than just numbers* This is because Camp, serlceus forages only during the day time whilst Camp*acvapimens is forages both day and night and this may account for the reason v?hy the foraging numbers differ* In terms of numbers and activity, these two species are among the dominant species of the savanna areas where the detailed study was carried out* 2 Camp*acvapiinensis and Camp.vestitus Both these ants nest in the soil but the existence between them is a passive association since Camp*vestltus prefers a no-shade area for nesting whilst Camp * acvapimensis was found to nest in areas with poor shade or broken shade or rarely in continuous shade* The former cannot be classified as dominant in terms of number of nests that were found in any area* In terms of foraging they were always very few and non-aggressive* The number of ants found in any given area was very insignificant* The ants Camp*vestitus do not compete with Camp * acvapimensis for space or food since they have different nesting habits and cannot exclude Camp * acvap imensis spatially or numerically. 3 Camp*acvapimensis and Annoma Driver ants of genus Annoma were found to nest mostly in no-shade areas and were found to be numerically abundant as well as dominant* Together with Camp♦ acvapimensis and Camp* sea?iceu3 they form the dominant species of the savanna ecosystem* - 40 - University of Ghana http://ugspace.ug.edu.gh Numerically however, these three species were many but the driver ants were more numerous than the rest and the most aggressive of them all. This is provably because these ants inflict very painful bites and easily bite as compared with the two other species that have been mentioned as timid, The driver ants were found nesting in some areas where Camp.acvapimensis was also found nesting but were never found in antagonism. Both ants had preference in nesting in the soil and their co-existence was probably due to the. avoidance reaction of Camp.acvapimensis and secondly due to their preferences for different shade regimes. Driver ants had preference for areas without shade whilst Camp ♦ aovapiaensis- has preference for areas with varying degmflfc of shade and are not many in shadeless areas. Numerical abundance In terms of numbers, driver ants are numerically dominant as well as aggressive. These ants forage both day and night but unlike Camp.acvapimensis« they move out of their nests in large numbers. In terms of aggression and numerical abundance, this group forms the dominant group of species of the savanna ecosystem followed by Camp.acvapirnensis.and then Camp.serioeus in that order. 4*4#5*4 Camp.acvapimensis and Platsrthyreus tarsartus Platythyreus species were not very common but were nesting in some areas that Camp, acvapimensis was found nest in/% University of Ghana http://ugspace.ug.edu.gh The aggressive nature of PI a t ythyr eus m s not observed and this is probably due to the timid nature of Camp . acvap imen sis which was found moving away from areas where any ant that is not of the same species was found. On a few rainy days however, a few ants, some of being workers of Camp, acvapimensis were found being carried away by the ants. University of Ghana http://ugspace.ug.edu.gh C H A P T E R F I V E N E S T A N D N E S T I N G H A B I T S University of Ghana http://ugspace.ug.edu.gh 5.1 INTRODUCTION Like all social insects, ant communities costruct nest f©r concealing their brood which is tended by the workers within the nests# Nests of such ants are found in different areas and may occur on leaves, wood or soil# Preference of nesting sites varies with the species. Crematogaster africana Mayr was reported constructing carton nests attached to the trunks of main stems or branch unions (Entwistle 1972), and Room (1 971) also reported that the same species nests in the canopy of cocoa trees. This indicates that the same species may have different nesting sites in the same area. Some ants such as Oecophylla longinoda Latreille are reported as founding nests in the cocoa canopies and using leaves for nesting(Leston 1968). Room(1971) also recorded that the same ant nests in the canopy of trees. Ackonor(1 977) reported the Catalaucus guineensis F. Smith may nest in dead portions of living trees where termites have already made holes or dead remains of the butress of forest trees where the termites had made the holes. In his work, Ackonor (1977) discussed in detail the nature of the nests constructed by the ants that he had studied. Very little work exists on the ants that nest in the soil. Out of some 157 species of ants collected by Room(1 971) in a cocoa farm, only 3 were reported at a depth of 30 cm below the litter on the soil. These were Solenopsis and Leptanilla - 43 - SHATTER 5 University of Ghana http://ugspace.ug.edu.gh species* ^ot much is known of the species that nest in the soil* Room(l 971 ) reported that Gamp* acvapimensis was found nesting in dead wood in the soil but did not investigate further when he reported that the same species were found foraging in the top 10 cm* of the surface litter and soil* Evans(1.971) reported that these ants Gfarp* acvapimensis were found carrying soil to construct tents over mealybugs which they tend. This finding wa.s in connection with his work on the black pod disease* On the same genus Gamponotus, Wheeler (1 922) reported that the group was mostly found nesting under stones, living in the soil, craters, under bark of trees and in hollow twigs. At Bolenzi, in the Republic of Gongo, Wheeler (7 922) found Cgjnp. acvapimensis nesting in the trunks of oil palms and at Faradje in the same country, the workers of the sar e species of ant were found attending plant lice on young leaves of orange trees. The reports of the various workers indicate that 2 types of nests are constructed by the species but in all cases no details of the nature of the nests are available* The castes found within each of such nests were not indicated and the report mainly deals with activity pattern of the species* On site preference, Majer (1 ?72) repo­ rted that the ants were found nesting in insola/ted ground* No information is available on the caste composition in various nests. Wheeler (1922) however, mentions that the ants of the genus Camponotus, form populous colonies and he - 44 - University of Ghana http://ugspace.ug.edu.gh recognised 2 forms ©f workers which he referred to as maxima and minima workers. Others which he mentioned were the alates males, females, and the pupae. No information on the phenology of the castes is available. In the following section, attempts were made to study the changes that occur in the nests during different months of the year and the effects of seasonal changes on the populations density of the ants. Composition of the different castes was also studied and their percentages determined. The nests were examined to ascertain the arrangement of the castes within the nests as well as the storage of food items in the nests. Caste ratio was determined at 3 months intervals by digging a number of nests. 5.2 METHODS A survey was made and the nests to be excavated were marked by the pegs pushed into the soil near the entrance of the nest. Seventy nests were marked outside the study area -for excavation. The reasons for locating the nests outside the study plots was to ensure that changes in the number of nests caused by the different seasons could be recorded on the study plots of changes in the populations of the ants, for a period of 12 months. Architectural specialities found within the nests were also recorded. Arrangement of brood within the nests was noted. - 45 - University of Ghana http://ugspace.ug.edu.gh 3?he excavation was timed to start at 12 noon on a predetermined day# This time was selected because of the observations made during the study of activity pattern which indicated that few or no ants came out of the nests just before and some time after noon. The ants were thus found to be less active and very few left the nests for foraging. The nests marked for excavation were the primary nests which were established in the soil. A close study of such nests was done for a period of four days and during this time all the secondary nefits used by the workers of the nest in question were marked so as to ensure that during the excavation period almost all the ants that constitute the caste of a particular nest were captured and their numbers recorded. Two nefcts were excavated each month. In all cases, excavation was done at 2 - week intervals (middle of the month and end of month). 5#2#1 Sampling Proceedure At the time of sampling a firm nylon thread was gently pushed into the nest as far as possible. Excavation was done using pick-axe, hoe, cutlass and soil chisel. Digging was done slowly in order to enable the worker to follow the thread and the tunnel into the chambers of the nest. To start, a pick-axe was used to loosen the hard surface crust and the soil was then collected from the entrance with the nylon thread acting as a guide into the inner chambers of the nest. When the lower areas were reached, cutlass and other implements were used for slow digging. - 46 - University of Ghana http://ugspace.ug.edu.gh This was done to prevent destruction of the ants in conccal­ led chambers. When a chamber was rea,ched, the contents we re collected into bottles containing 20/& alcohol. These containers were placed very close to the nest so that transfer of the captured ants into them could be done quickly and with ease. All the captured ants were put into the same container. When more than one tunnel was seen leading into different underground chambers, a piece of wood or stick was used to block the entrance to one of them and one tunnel was dug at a time. When digging was complete in one direction and the end of the tunnel was reached, the other blocked tunnel was dug to Gollect the contents of the chambers that they led into. Workers and soldiers tending aphids and scale insects were captured and added to the castes. No records were made of the number of ants in secondary nests. Any ant of the same species that came searching into the nesting area with the character­ istic raising of the gaster was captured and added to the caste. 5*2.2 Sorting Sorting was done a day or two after excavation of the nest. When the population in any nest was too large, sorting took one or two days to complete but with very small popula­ tions, sorting was completed in only a day. The ants that had been sorted were kept in different containers in 5OJ6 alcohol and the different stages of the castes being the University of Ghana http://ugspace.ug.edu.gh larvae, pupae, and adults were preserved separately,# The ants were also sorted out into workers and soldiers* The sorted ants were kept in a cupboard for counting* Guests of the nests were only recorded as present or absent but their exact numbers were not determined* 2Food items found in the nests were kept in 20J{. alcohol and analysed later in order to determine the nature of the food the ants take to their nests* 5*2.3 Counting Actual counting was done for each of the different castes, of the brood. The ants were picked from one container, their numbers recorded and they were then placed in another container. Counting of the larvae took a long time and in cases where many larvae were found, they were counted with the aid of a hand lens. The ants were recorded as soldiers or workers but it was not easy to distinguish between the pupae that were developing into soldiers or workers. It was however, easier to sort the pupae and larvae of the alates since they were rather large as compared to the larvae and pupae of the worker and soldier castes. 5*2.4 Head measurements During the sampling period, head measurements were taken for 50 workers and 50 soldiers or more, found in a particular nest. After a time, the measurement of the worker - 48 - University of Ghana http://ugspace.ug.edu.gh caste was done after two months intervals as they were not found to differ significantly from other workers in other nest s.Measurements for the soldier caste continued because the soldiers were observed to be of two types. One type of them was seen to be foragers and collectors of food whilst the second group was found to only guard the workers and other food collectors. The head widths of most of the soldiers were the same so the measurement took into account the length rather than the width. By means of an eye-piece micrometer, the head measurements were taken and the difference between foragers and guarding ants found in the different colonies was determined. The importance attached to finding foragers among the soldiers was because the brood in some 0f the nests was very large for the workers alone to feed them and it was noted that the soldier caste were also foraging during the period when the brood increased in the nests. Head measurements of the soldiers in other nests were teken to find out if any significant difference existed between the soldier caste of the different nests. No measurements were done for the alates as their numbers were very few during the study period. .5 The Nest The tunnels that formed the entire nest were measured with a thread as digging went on. At the end of the digging, the longest thread used for the measurement of the nest University of Ghana http://ugspace.ug.edu.gh tunnels was teken and the results recorded && che actual depth of the nest* Due to the inclinations of most of the nests, two depths were considered, (a) Vertical depth: this depth was measured at the end of digging. The last chamber of the ants that was considered as tne deepest of the entire nest was measured with the ground level as the starting- point. The greatest distance of the nest from the surface of the soil was then obtained and the results were then recorded! (b) Actual depth: this depth was measured by using a thread. Prom the entrance of the nest the tunnel was followed and when the contents had been collected, the thread was used to line the centre of the nest until the end of excavation was reached* The digging continued and at the end of collecting the contents, the thread was used to take the measurement of the different tunnels until the total depth of the deepest nest was found* The thread was then measured with a meter - rule* 5«2.6.Type of soil in which nesting took place Observation during the search for nests revealed that the nests were found in certain areas and this stimulated investigation for the soil type in which the nests were mostly found. For this purpose, a sample of the soil in which the nest was found was taken and put into a measuring cylinder* The volume of the soil was found out and recorded* - 50 - University of Ghana http://ugspace.ug.edu.gh Water was poured into the soil and it was then vigorously shaken for all the particles to be loose and dissolve if possible. The cylinder was then put down for the particles to settle down. The sides of the container were washed down sind the whole experiment allowed to remain for about four hours. The different soil percentages were then classified as mixture of clay and sand, sand only, silt and clay or silt and sand, or clay and litter. This classi­ fication ia ';hen related to abundance of nests within the given area and also considered as the basis of preference for nesting areas. 5*2.7 Position of nest entrance in relation to vegetation Area with nests and thick vegetational cover were cleared to determine the effects of increased temperature places and low humidity on the nesting habits of ants- without nests were provided with artificial shading over a period of time (t month) to find out if the ants would construct nests in these areas due to the shade that had been provided in the area. Nests in the roots of grasses were carefully studied and the foliage leaves of such grasses removed very close to the roots so that the effects of such environmental factors as rainfall, light intensity, evaporation, low humidity and temperature fluctuations may affect the ants and the be­ haviour of the ants recorded. Thirty such experiments were performed. - 51 - University of Ghana http://ugspace.ug.edu.gh The foliage leaves; were never allowed to grow more than 1! centimeter above the height to which it was trimmed at the beggining of the experiment, Effects of flooding on the nests Since almost all the primary neets were found in the soil, observations were made on the nests just before a rainfall and immediately after a rainfall* The conditions in which the entrances of such nests were fou^d were recorded and any peculiarities also scrutinized. Just after the rainfall, such nests were observed and the condition under which the nest occurred was recorded. During the dry season, water hose was also used to flood the nests to find out the behaviour of the ants during the flooding period. Type of nests Attempts were made to find out the types of nests constructed by the ants and these were then classified as primary or secondary nests. The occurence of such nests has been pointed out earlier (Chapter 4)* Observations/Results Contents of the nests: The contents of the nests are made up of larvae, pupae, workers, soldier and few reproductives during some periods of the year. University of Ghana http://ugspace.ug.edu.gh The foliage leaves were never allowed to grow more than | centimeter above the height to which it was trimmed at the beggining of the experiment. Bffects of flooding on the nests Since almost all the primary nests were found in the soil, observations were made on the nests just before a rainfall and immediately after a rainfall. The conditions in which the entrances of such nests were found were recorded and any peculiarities also scrutinized. Just after the rainfall, such nests were observed and the condition under which the nest occurred was recorded. During the dry season, water hose m s also used to flood the nests to find out the behaviour of the ants during the flooding period. Type of nes-ts Attempts were made to find out the types of nests constructed by the ants and these were then classified as primary or secondary nests. The occurence of such nests has been pointed out earlier (Chapter 4)* Observations/Kesults Contents of the nests The contents of the nests are made up of larvae, pupae, workers, soldier and few reproductives during some periods of the year. University of Ghana http://ugspace.ug.edu.gh (a) Larvae: The larvae were of different sizes and the larvae of 0ach of the castes were found to be of different stages of development. The small ones were very tiny and could be seen only by the use of a hand lens. By the use of even the binocular microscope, no eggs were found in any of the nests* Some of the larvae were very tiny and could easily be taken for eggs. The larvae found show clear segmentation but the segmentation became very marked as the larvae moulted* By comparison, the larvae of the ants that were younger appeared slightly roundish and whitish and became creamy as they became older. Each larva bears distinct hairs on the body and they become more pronounced as the larvae become larger and older. There were no visible head appendages on the larvae. The larvae of the different cashes could be distinguished when they became mature and the soldiers appeared larger (fig.9b) then the workers (fig.9c) and the reproductives were the largest (fig*9a) in the overall consi­ deration* a. result, three different larval types could be distinguished by the differences in their sizes. In some cases it was not easy to determine what a larva would develop \i\tn as very young stages of soldiers and workers looJt alike. In such oases, the presence of hairs was used to mark the differences. (*0 Pupae: The pupae of C.acvapimensis are enclosed in cocoons. The pupa® were distinguishable when size alone was considered. The worker caste pupae were very small sized - 53 - University of Ghana http://ugspace.ug.edu.gh (a) Larvae: The larvae were of different sizes and the larvae of each of the castes were found to be of different stages of development. The small ones were very tiny and could be seen only by the use of a hand lens. By the use of even the binocular microscope, no eggs were found in any of the nests* Some of the larvae were very tiny and could easily be taken for eggs. The larvae found show clear segmentation but the segmentation became very marked as the larvae moulted. By comparison, the l a r va e of the ants that were younger appeared slightly roundish and whitish and became creamy as they became older. Each larva bears distinct hairs on the body and they become more pronounced as the larvae become larger and older. There were no visible head appendages on the larvae. The larvae of the different caaebes could be distinguished when they became mature and the soldiers appeared larger (fig.9b) then the workers (fig#9c) and the reproductives were the largest (fig#9a) in the overall consi­ deration. a. result, three different larval types could be distinguished by the differences in their sizes. In some cases it was not easy to determine what a larva would develop iftiact as very young stages of soldiers and workers loot, alike* In such oases, the presence of hairs was used to mark the differences. (b) Pupae: The pupae of C*acvapimensis are enclosed in cocoons. The pupae were distinguishable when size alone was considered. The worker caste pupae were very small sized - 53 - University of Ghana http://ugspace.ug.edu.gh University of Ghana http://ugspace.ug.edu.gh - 54 - and two types of cocoons were distinguished* The first type of worker caste were smaller forms and the second type were smaller but slightly enlongated in appearance (Fig.lOa). The mandibles which could be seen through the cocoon were structurally smaller as compared to those of the soldier caste. The soldier caste had slightly larger and longer cocoons (Fig.lOb). The two types of adult soldier castes could not be distinguished during the pupal stage of their development. The alates had very large cocoons but the head and mandibles were very small in size (fig. 10c) but could be distinguished from other members of the caste. In all pupae it was observed that the mandibles were very prominent in the cocoon. The mandibles of the soldier caste were very exceptionally prominent and the size of the mandibles could easily be used to distinguish the soldiers from the larger pupae (workers). (c) The workers; The workers are small in size as compared with the rest of the castes. The eyes of the ants are very small and are situated behind the middle of the head. The head is narrower in front and broader in the middle. The thorax, fits into a grooved depression in the head at the posterior side of the head. The eyes are small and the antennae are 12 - jointed and its scape is situated a little farther from the end of the clypeus. University of Ghana http://ugspace.ug.edu.gh University of Ghana http://ugspace.ug.edu.gh The mandibles are la,rge and triangular with the teeth found as coarse projections on the bfOAd surface. The thorax is broad in front and is laterally compressed towards the poste­ rior end. The gaster of the ant is slightly larger than the rest of the body and found to be raised in excited ants. The legs are long and well developed. Bach tibia possesses a spur just before the tarsus (Fig* 1l)« (d) The soldiers: Morphology of the soldiers (Fig. 12) is the same as of worker caste but differs in head size which is broader as well aa> longer than the worker caste. Two types of soldiers may be distinguished by head size. The first type has broad and short heads and the£e were earlier referred as the foraging soldiers (Fig. 13a)# Their mandibles are less coarse as compared with the man­ dibles of the other soldiers. The mandibles even though triangular were smaller as compared to the mandibles of the other soldiers. The second type of soldiers (Fig. 13b) were earlier ref^ped to as the guarding soldiers during foraging. These soldiers had broad and long heads and their mandibles were very large and triangular with coarse teeth on the broad surface of the mandibles. By comparison, their heads appeared narrower than those referred to as the foraging soldiers. (e) The females: The females are the reproductives of the caste. Their head structures appear like the worker caste - 55 - University of Ghana http://ugspace.ug.edu.gh 0 = O CE L LUS FIG'. 11 -.Worker of Q- ac vapimensis X'250 University of Ghana http://ugspace.ug.edu.gh FIG:12: Soldier of C. acvapimensis X 120 University of Ghana http://ugspace.ug.edu.gh FIG: 12: Soldier of C. acvapimensis X 120 University of Ghana http://ugspace.ug.edu.gh (a) forager ( b ) guard FIG.'13-Heads of the two kinds of C- acvapimensis soldiers, X2 50 University of Ghana http://ugspace.ug.edu.gh - 5 6 - but oould be easily distinguished by the presence of wings as well as their very large gaster s. They have ocelli and the wings are very long. (Fig. 14)* (f) The males: THe males are smaller and slender with short heads. Their eyes appear more prominent and ocelli are also present. Their gasters are comparatively elongate, legs are very slender (Fig* 15) • (&) The queens: The queen of the nest is just like the female of the caste but differs only in the absence of the wings (Fig# 16)# 5*3*2 Guests of G« acvapimensis During excavations, the only foreign insects in the nest of the ants appeared to be termites Microtermes spp. Investigation revealed that the nests of these two insects were separated by thin walls and these walls were broken during excavation when the ants attacked the termites and carried them away. In the nests, any termite present was dead and was among a heep of food items that the ants had collected into the nest and could not be considered as the guests of the ants. Those nests that were found in the wood, the termites were observed to have left the nest before the ants might have occupied the nest at a later stage. University of Ghana http://ugspace.ug.edu.gh University of Ghana http://ugspace.ug.edu.gh o=oceUus FlG'16: De-aLa te queen of C. acyan ixnen .s i^ x 120 University of Ghana http://ugspace.ug.edu.gh o=ocellus RG.’16 : D e -a la te queen of Q- x 120 University of Ghana http://ugspace.ug.edu.gh 5»3*3 Phenology of brood and castes During the study period, the brood and adults in the excavated nests were counted and the results are given in table 14* Two papulation peajks occurred between mid - February and the end of March# The largest population was 1 ,572 per nest while the smallest was 227 recorded during the middle of January* On the whole, an average of 643 indi­ viduals per nest was obtained* Further information on the phenology of the brood and castes is given in Figs* 7 and 8* The larvae were observed to have fluctuating peaks with three of them distinctly marked* The highest peaks occured towards the end of December, middle of April and the middle of July* ^he lowest peaks occurred at the middle of January and during May* The percentage of pupae per nest followed a pattern opposite to that of the larvae in that during the periods when the larvae were more, the pupae were few and vice versa. Differences occurred in the middle of November, and the middle of December, 1976 and the end of May, 1977 when both larvae and pupae had lower peaks* The highest peaks of the pupae occurred towards the end of September and the minor peaks at the end of February, April - middle of May* The percentage of soldiers present was almost constant except in the middle of November when a high peak occurred and in the middle of March when their numbers declined to a negligible percentage (almost zero)* - 57 - University of Ghana http://ugspace.ug.edu.gh FI G .’ 7 Ph en ol og y of br oo d in va rio us n es ts CD C^* (Ti * L A R V A E 19 77 • • P U P A E TI ME ( M O N T H S ) University of Ghana http://ugspace.ug.edu.gh University of Ghana http://ugspace.ug.edu.gh Table 13: Composition of the castes of the various nests CamT)>acvaplmensis excavated during the study period* Date larvae Pupae Soldiers Workers Winged Total 30/ 9/76 52 197 94 164 0 507 15/10/76 157 179 120 216 0 672 30/10/76 62 108 73 79 0 672 15/11/76 68 29 82 104 0 282 30/11/76 42 152 52 170 0 416 15/12/76 68 170 69 241 0 548 30/12/7$ 293 192 62 217 18 782 15/ 1/77 9 67 38 111 1 227 30/ 1/77 114 124 43 166 3 451 15/ 2/77 415 558 108 323 0 1405 28/ 2/77 320 789 103 337 22 1572 15/ 3/77 169 416 81 381 0 1048 30/ 3/77 225 334 62 27 6 0 898 15/ 4/77 220 51 54 221 13 560 30/ 4/77 51 125 20 68 0 265 15/ 5/77 34 241 56 187 5 532 30/ 5/77 18 35 61 204 20 338 15/ 6/77 219 199 131 361 6 916 30/ 6/77 134 109 69 124 9 445 15/ 7/77 320 79 101 316 23 839 30/ 7/77 194 121 62 211 2 590 15/ 8/77 214 201 93 179 6 693 30/8/77 120 172 79 133 0 504 15/ 9/77 94 214 109 £92 18 697 University of Ghana http://ugspace.ug.edu.gh recorded. This gave an average of J6 soldiers per nest# 5*3*4 Head measurements The head measurements for the workers and the soldiers are given in appendix tables 2-13. The maximum of the soldier (from mandibles to where the head is fixed to the thorax) was 2.50mm and the minimum 1.39mm. There were 2 extremes! (a) short heads and (b) long heads. It was also found out that those of the former category had broader heads and those of the latter group had narrow heads (Pigs. 13a and 13b). These differences were found in the soldiers from most of the nests. (Pigs 1 7 - 2 2 are the results of the head measurements found in the different ranges). Fgom the histograms it is clear that two groups of soldiers are present! (i) those with broad but short heads and (ii) those with narrow and long heads. There were no intermediates between these extremes but in certain colonies (Pigs 17 and 18 ), only one group of short head soldiers were found while in other nest (Pigs 19 -22), the range of head length shows 2 groups with Pigs 1 9 22 showing a. wider range for the head lengths. This indicates that in Pigs 17 and. 18, there is only one group of soldiers. In the worker caste, theafcwas no difference in all the colonies. The head range was found to lie between 1.13 and 1 .72mm but most workers tended to fall in the middle group (Pigs. 23 - 28). - 59 - University of Ghana http://ugspace.ug.edu.gh CD I— [/)UJ Z SI 6 lL O z LU QsX in InUJ z t N 0 LL ”T“<2) a a.«ta a*oft HEAD LENGTH (m m ) University of Ghana http://ugspace.ug.edu.gh tCO V— m W z £ r "~ z CM T U3 T C4 00 T~ »* 00 0) UO Lp V— UJ 2 in ^ cm 0) fS 00cr o ^ cs ~rCD T “ CM 00 ~r A 0 N a n n 3 w y ni t >m University of Ghana http://ugspace.ug.edu.gh 00 r* h- l/) UJ z: a o Ee o z LU Q a X in in iu z N 0 IL r ' ~ J> **1, LU"* UJ — Q 0 acvapimensist a savanna ant constructs nests as deep as 63*1 cm# (b) Actual depth of last cell This consideration was not found to be benefiting as some ants were found constructing nests with cells a few milli­ metres within the soil and the cells were not distinct from each other but very shallow* The number of chambers that constituted the nest was therefore considered and it was found. Soils in which nesting took place V6ry few nests were founding in the sandy area and this might be due to the poor nature of the soil as the ants were found to be nesting near food sources* ~ 70 - University of Ghana http://ugspace.ug.edu.gh It is considered that under very favourable consitions the ants might be found nesting in the sandy soils in areas that have good cover of vegetation. Effects of flooding Flooding seems to have little or no effect on the ants apart from drowning some of them. On the whole the ants seem to have no nesting preference in relation to flooding* Nest cleaning Nest cleaning seemed a constant day-to-day activity but it also took place after the rains. After the rains it became necessary as the materials that had entered the nests were brought out. This activity was extended to include nest expansion, nearly all nest cleanings tend to include nest expansion# Types of nests and arrangement of brood The present work has shown that there may be more than one nest and hence there is always one primary nest found in the soil or wood and many secondary nests. This tends to agree with the findings of Wheeler (1922) who found the species nesting in the trunks of oil plam and some of the workers also tending scale insects and aphids on orange trees. It is evident that the nests in which they were found tending the insects remained as nests as long as those insects kept feeding on the plants. - 71 - University of Ghana http://ugspace.ug.edu.gh C H A P T E R S I X ACTIVITY PATTERN OF CANPONOTU3 ACVAPIII3TSIS University of Ghana http://ugspace.ug.edu.gh - 72 - ACTIVITY PATTERN OF CAJ.TONQTUq ACYAPHl^TSIo INTRODUCTION Like all social insects, ant communities contain a number of castes which perform different types of work. The time at which their duties are performed and the duration constitutes activity pattern of the community* The main activities include the building of nests, foraging, mating, feeding of the brood and killing of prey* The time or period during which the ants emerge and begin their activity is influenced by environmental factors* Wheeler (1930) reported that the ant Prenolepis icrparis Say begins to emerge as soon as the tempera tare reaches above freezing point, increasing in number as the temperature rises until the temperature reaches a peak between 45° and 65°F* However, as the temperature rises above 85°F, their numbers decline* Tabolt (1943) also confirmed this report and added 'kha'fc Prenolepis imparis were found to react to temperature but to a lesser degree* Ayre (1958) however, working on Formica subnitens Creighton, concluded in his findings that relative humidity was of a little importance in limiting ant activity* Ayre (1958) attibuted the effects of tempera­ ture to the correlation between light, temperature and humidity* Ackonor (1977) working on Cataulaucus /?uineensis F* Smith reported that air temperature and relative humidity CHAPTER SIX. University of Ghana http://ugspace.ug.edu.gh also seem to be important factors influencing ant activity but felt that their effects were indirect whilst light acted directly* He further stated that light intensity was the most important single factor directly or indirectly* However, environmental factors other than temperature and humidity may also affect the activity pattern of ants* Ayre (1958) further reported on Formica subnitens that on cool days the ants had one foraging activity but on warm days the ants had two foraging activities, 'Phis tends to indicate that temperature had effects on the foraging acti­ vity of the ants. Rainfall was also another factor that affected the activity pattern of the ants. Ackonor (1977) reported that Gat.guineensis stopped their foraging activity during rainfall which led to a delay in commencement of activity. This he noted also led to a delay in the start of activity on the next day* Activity pattern can also be affected by the brood present in the nest at any given time* Schneirla (1944) pointed out that the type of brood present in the nest (larvae or pupae) influenced the activity pattern of ants. Yowles (1955) reported that temperature and humidity alone could not account for the differences found in the activity pattern of the ants because the ants seemed to forage despite some unfavourable temperature and humidity effects. He observed that the effect was more the cause of the presence of the brood (eggs, larvae, and pupae) rather than - 75 ~ University of Ghana http://ugspace.ug.edu.gh the effects of temperature and humidity. During activity pattern, ants tend to e-hibit all sort of behavioural patterns and Vowles (1955) attibuted all these to nervous excitation caused by factors such as hunger and availability of food in the environment. Adabie (1977) attributed the stimulation of foragers in the nest of Cfrematogaster depressa (Latreille) to forage to the returning of foragers into the nest and the probable scattering of the developing larvae in the lower chambers of the nest. She felt that the returning foragers entered these chambers and stimulated more workers to forage and increase food supply. In the present study, investigations into the activity pattern of Camp.ac vapiaen sis were carried out. Several factors were considered. These included the effects of the seasons, the effects of such environmental factors as humidity, temperature and light intensity. Other factors that possibly influenced the activity pattern and foraging of the ants wefe studied. These included the presence of brood and food in the environment. - 74 - University of Ghana http://ugspace.ug.edu.gh MATERIALS AIID M^iTIODS Foraging activity In the study of activity pattern of Gamp. acvapimensis two nests were selected for 12 months of observations. This was done to ensure that the effects of any changes in populations could be noticed. One nest was selected in the grassland area of Plot A and the other in the farm land area of Plot B. The two nests were selected for study in order to investigate the effects of seasonal changes on the activity of the ants as well as the effects of farming on the activity of the ants. In all cases the numbers of ants crossing the census points into and out of the nests were tallied on a card and later recorded in a field notebook. It is to be noted that there is only one opening to the nest and the ants use this as the exit and entrance at the same time. It was therefore easy to tally the numbers leaving and entering the nest at the same time. Uumbers of ants entering the nest with food were also recorded. 6,2.1 *1 (a) Diurnal activity pattern During the study period the number of ants that entered and left the nest as well as those entering the nest with food were recorded as explained above during the first ten minutes of every hour (i.e. 6.00 - 6.10 a.m. etc). The behaviour of the ants such as moving In trails, or fol­ lowing others in search of food was carefully studied as - 75 - University of Ghana http://ugspace.ug.edu.gh well as "kke effects of certain environmental factors such as relative humidity of air, temperature of the air and nest# Other factors like clouds cover and mist were also taken into consideration. The humidity of air was measured with a hygrometer and the temperatures of the air and nest were recorded with a Grant* s temperature recording instrument (Grant1 s miniature recorder). The instrument was set to record the temperature at hourly intervals. The temperature of the nest was measured by pushing the sensitive end of the cable into the nest through the entrance deep down and at the same time ensuring that the nest was not blocked by the cable. To obtain accurate results, the cable was pulled out gently and cleaned about 2 days to the time for study of activity pattern and gently pushed dovzn again. This was done to remove any deposition of soil on the sensitive part of the cable and rendering it ineffective. Two days were considered sufficient to allow the ants to get used to the presence of the cable which had been introduced. The relative humidity of the nests was measured at specific hours (e.g. 6a.mf 7 a.m. etc), but the activity was studied over a period of ten minutes. °*2.1.2 Nocturnal activity pattern The ants were observed to be active both day and night and therefore it became necessary to consider the night activity as well. This was carried out using a hurricane lamp near the entrance of the nest. The lamp was covered - 76 - University of Ghana http://ugspace.ug.edu.gh w ith a red cellophane and was used throughout the period o f study# Other colours tried had certain disadvantages and were discontinued, 5*2*2 Foraging space and distance In trying to determine how far the ^nts moved from their nests when they went out to forage, eosin powder was used to dust the ants and then they wore later followed as they went out* On specific days, dusting of the ants was done for about 2. hours and the ants were then followed into the field as well as on vegetation* r£he farthest point that they moved to was recorded* *flien it “became difficult to determine the identity of the marked ants, a drop of water was put on the gaster of such ants* All ants that were dusted showed a characteristic eosin colour on contact with water. Other methods used to study the distances that the ants moved from their nests was to dust ants foraging on leaves and disturbing them. Such ants moved away towards their nests and the distances from their nests to such foraging areas were measured* Such investigations were possible only during the day as the ants could not be tracked easily in the night. It became necessary therefore to use a suitable method for the distance that the ants moved during the night. Since the ants are fond of sugar, sugar baits were used for this purpose. In order to be sure if the ants were from the nests that were being consi­ dered, eosin was dusted on the ants that found the baits - 77 ~ University of Ghana http://ugspace.ug.edu.gh and then th e entrance to their nests was guarded w ith a h u rr ic a n e lamp, observing the ants that entered the n e s t. Any ant that entei^ ed the nest was examined with a drop o r wa ter as it passed over a blotting paper. Presence of eosin showed up on the blotting paper. An attempt was made to find out the differences in foraging distances during the dry and wet seasons. •This was considered necessary as the green leaves on which the ants were found foraging dried up in the dry season and the ants were found to travel long distances in search of :green leaves. To obtain accurate results on foraging behaviour of the smaller and larger workers, these were followed differently onto the vegetation on which they foraged and their activities observed. Observations were also made on the ants that brought food into the nests. 6*2.3 Induced foraging Since the ants were found to forage day and night and the only interval with least activity was during mid-day when li^it intensity and temperature were very high, other factors that caused the activity were also investigated. 6.2*4 Ants - phytophagous insect- associationship The activities of the ants in relation to attendance of plant sap-feeders we re studied. This took into consideration the type of associationship that existed between the ants and the pests. For this purpose, a search for secondary nests or tents was carried out .and the nests found with pests - 78 - University of Ghana http://ugspace.ug.edu.gh - 79 - opened and the contents collected into 70/i alcohol for classification. The association between the ants and the sap-feeders was then classified as obligate or facultative. In this classification, the presence or absence of tents and other protection afforded by the .ants was taken Into consideration. 6*2.5 Flight activity Flight activity of the ants was studied using a light trap for a period of 13 months. C&tdies were collected and the number of alates found recorded. The number of alates cau^it over a period of one month was added together. Pood items collected by ants On days that foraging distance was determined, the plants with foragers were noted and identified. The specific activities of the ants on such plants were recorded. The plants were then grouped under different categories such ass (a) plants on which ants foraged daily, (b) plants from where the ants collected food materials and (c) plants on which ants tended sap-feeders. The food items collected by the ants were seized and analysed. Plants on which the ants were found to be moving on their foraging errands were not recorded but were considered as plants on their way* During such recordings, any damage that the ants caused to the plants was also noted. To investigate further the type of association existing between the sap—feeders and the ants, University of Ghana http://ugspace.ug.edu.gh the ants were sprayed off at the bases of such plants or grease was applied around the bases to prevent the ants from visiting the sap-feeders and observing the effects of this on the pests* ;#3 OBSSRVATIOIIS/FuJSITLTS ;,3*1 Fora-yin.? activity of Gamp, acvapifflensis A careful study revealed that the activities of the ants followed the sane pattern and rythmn on plot A and B in the wet season but slight differences were found in the dry season* The differences were the resalts of two major factors, drought which caused the leaves of the plants to dry up and harmattan which increased the evaporation of water from the ants. On the whole, the ants wer-? observed to be active both day and niiht* The periods varied and could be classified as a period with least activity or a period with maxijpm activity. Both soldiers and workers were observed to be active all day and night but the workers were observed to be more active than the soldiers p r o b a b l y "because of their numbers and secondly because of the functions of the soldiers in the nest. The soldiers came out to forage but they were rarely found coming out alone* They mostly followed the workers. In most cases they returned to nests alone or with the workers* The activity pattern of the ants was then classified as diurnal or nocturnal, depending on the time of activity. - 80 - University of Ghana http://ugspace.ug.edu.gh 6*3*1.1 activity pattern The general trend of activity each day was £ueh that some workers left the nests while others entered them but in the mornings, (6 a#m#) fewer ants were found crossing the census point* Most ants that crossed the census point were leaving the nest whilst only a few were entering. On any ordinary day, (i.e. days without rain or any environmental factors causing changes in the climatic and soil conditions), the number of ants found leaving the nest and those entering indicated that two peaJcs of activity occurred# Between 8 a#m. - 11 a#m, the ants were observed to be active and their activity ceased or became low between 11 a#m - 2 p#m# The same type of activity pattern was observed from 2 p.m - 4 P*m* This resulted in the two peaks of activity foxind in the ants. One of the activities occurred in the morning and the other in the afternoons (figures 40 and 41 )• On these days, the activities were found to be influenced by such environmental factors as ligii intensity, temperature and humidity but the light intensity, seemed to :affect the‘ other two factors within the environment On days when the sun rose earlier and the light intensity caused a quick rise in temperature and lowering of humidity, the ants were inactive even though certain factors such as the presence of brood could contribute to an increase in the activity# On such days, very few ants left the nest between 8 a#m and 11 a#m when in fact activity came to a standstill - 81 University of Ghana http://ugspace.ug.edu.gh A N T S ■ f ig : 32 TI ME (HRS) TE MP . CC ) Vo RE L. HU MI DI TY University of Ghana http://ugspace.ug.edu.gh F IG : 33 TIME (HRS) TE M R( *C )/ ‘^ RE L- H U M ID IT Y University of Ghana http://ugspace.ug.edu.gh FIG : 34 TE M P (# C ) /* /. R E L, H U M ID IT Y University of Ghana http://ugspace.ug.edu.gh FIG: 35 TIME (HRS) T EM R ( C ) /* /o R EL . H U M ID IT Y University of Ghana http://ugspace.ug.edu.gh FIG: 36 T IM E (.HRS) TE M P. (C ) I *to R EL . H U M ID IT Y University of Ghana http://ugspace.ug.edu.gh NO - OF AN TS FI G: 37 TI ME (HRS) TE MP . C C ) I *7. RE L. HU MI DI TY University of Ghana http://ugspace.ug.edu.gh F| g: 38 TIME CHR5) TE M P .C C ) / */. R EL . HU M ID IT Y University of Ghana http://ugspace.ug.edu.gh f i g: 39 TIME (HRS) T E M P . C C ) / 7o RE L. H U M I D I T Y University of Ghana http://ugspace.ug.edu.gh f | g : 40 2 2 -4 -7 7 TIME (HRS ) TE M P. °(L /°/ o RE L. H U M ID IT Y University of Ghana http://ugspace.ug.edu.gh NO- OF A N TS f | U. 6 - 5 - 7 7 TIME (HRS) TE M P. °C / % RE L. HU M ID IT Y University of Ghana http://ugspace.ug.edu.gh FIG: 42 17-6-77 TIMEtHRS.) TE M P . °C ./° /o R E L. H U M ID IT Y University of Ghana http://ugspace.ug.edu.gh fig : A3 A l l d l W D H " 1 3 H ° /o / 3 „ d W 3 i T IM E (H R S University of Ghana http://ugspace.ug.edu.gh till about 4 p.m when the activity was resumed. The activi­ ties had two peaks but each wan so slow and the ants were not seen or were not active with very few just crossing the census point and returning immediately into the nest (figures 32 and 33)# On several other days (figures 35t36 and 38) j light intensity was high but the ants remained active all day, despite the high temperature and the low humidity (figs 35 and. 36) but a careful study of the possible factors indicated that the presence of brood in the nests was high during the same period and this was the probable cause of the increase in activity of the ants* Cloudy days seemed to induce different behaviour patterns as the ants were found to be active all day despite low tamperature and high humidity* (i) Dry season Activity in the dry season seemed to foliow the same rythmn as found on any ordinary days when it was sunny. The activity pat Gem of the ant did not differ from that found during the ordinary days as shown in figs 32 and 33. These days were characterised by a quick rise in temperature and consequent lowering of humidity (Figs 37 opd 3? ) • The number of ants leaving the nests was small in the dry periods. But the ants were1 active just before 8 a.m and just after 4 P*®* On several occasions, no ants were found leaving or entering the nest at all. This inactivity became very marked during the harmattan when ants were mostly confined - 82 - University of Ghana http://ugspace.ug.edu.gh to their nests during the mornings and afternoons. There seemed to he no activity at all during tne day (Fig 37 )• This as stated earlier affected the activity of the ants in the night* Temperature, humidity and ligfrt intensity also affected the ants hut this became marked a& the harmattan affected all these factors. On a particular day (25th February, 1977) the ants came out in their numbers despite the harmattan. At 6 a.m as many as 19 ants were found leaving the nest‘d and as many as 14 were carrying food into the nest. Trailing the ants to inhere they brought the food, the ants were found near a termitarium broken loose by a farmer from ^ich the ants were collecting their food. Food carries were many which is rather unusual and by 2 p.11, the number of ants, that were leaving the nest and those that were entering had doubled. As many as 36 ants entered the nest, 29 (81J0) of them carrying food. By 2 p.mf 88 workers and soldiers left the nest and 35 entered, 14 of them carrying food (Fig 38). Similarly, in Fig 43 5 the ants we re active all day but on this day, it was observed tha humidity was slightly higher and the activity was also high due to the presence of a food source. (ii) TTet season During the wet season, activity pattern was different in that rainfall became another environmental factor that affected the ants. On 11th March, 1977 j the day was cloudy at 10.00 a.m„ and a heavy rainfall followed around 12 noon. University of Ghana http://ugspace.ug.edu.gh The ants were not found coming out and none left the leaves into the nest but moved to the lower surface of the leaves. For a period of 2 hours of rain, the ants stayed out, keeping themselves under the leaves on which they were foraging. Just after the rain, the ants began moving towards the nest and others moved to the upper surface of the leaves to forage. The ants became active soon after this (figure 39). Figure 34 shows another day when rainfall in the night was followed by a quick rise in temperature and lowering of humidity due to high intensity of light in the early morning. The rain stopped by 6*30 a.m. in the morning but the ants became active the whole day until 4*00 p.m and their activity rose again after 6.00 p.m. 63. !•£ Mffht activity The night activity of CamT)eacva~Pimensis followed a different pattern from that of the day time. On most days, the activity pattern of the previous day continued late into the night and sometimes ended by 8.00 p*m (figs 44 - 48). The ants became active again at about 5 or 10 p.m. The period between midnight and 1 a.m was the time for nest cleaning. During this period, the ants were found bringing out pieces of sticks, straw, food particles and soil and depositing such items a few centimetres from the entrance of the nest. On days that the number of larvae and brood were found to be high in the nests, the ants were observed to be active all night. Peaks of such activities did not follow any rythmn - 84 - University of Ghana http://ugspace.ug.edu.gh FIG: 44 28-10-76 T E M P . C #C ) / °/ „R EL - H U M I D I T Y University of Ghana http://ugspace.ug.edu.gh FI G'. 45 11-11-76 University of Ghana http://ugspace.ug.edu.gh FI G!46 2 5 - 1 1 - 7 6 TE M P( °C .) / % r el h u m id it y University of Ghana http://ugspace.ug.edu.gh FIG: 47 23-12-76 TE M P .C C ) /% > RE L. H U M ID IT Y University of Ghana http://ugspace.ug.edu.gh FIG : 4 8 20-1 - 77 T IM E {H R S ) T E M P . ( t ) I */• R E L. H U M ID IT Y University of Ghana http://ugspace.ug.edu.gh N O - OF A N T S FI 6 :4 9 4-2-77 T IME (HRS) T E M P .C C ) / ’/ .R E L . H U M ID IT Y University of Ghana http://ugspace.ug.edu.gh NO . OF AN TS FIG: 50 IB-2-77 TI ME ( HRS ) TE M P. CC ) / 7p R EL . H U M ID IT Y University of Ghana http://ugspace.ug.edu.gh NO. OF AN TS •FI G: 51 4-3-77 T1 ME (HRS ) TE M P. C °C ) / •/. RE L HU M ID IT Y University of Ghana http://ugspace.ug.edu.gh NO . OF A N T S FI g: 52 3 0 -4 -7 7 Tl MECHRS) T E M P C C ) / */. RE L H U M I D I T Y University of Ghana http://ugspace.ug.edu.gh FIG: 53 14-5-77 T IM E (HRS) T E M P r C ) / °U RE L. HU M ID IT Y University of Ghana http://ugspace.ug.edu.gh FIG: 54 28 -5 -77 TI ME CHRS) ' TE M P- l *C ) / c/o RE L H U M ID IT Y University of Ghana http://ugspace.ug.edu.gh (figs 47,' 49, 50 and 51)* On some such days (figs 47 and 51), as many as 4 peaks of activity were observed and they did not follow any rythmn. On certain days only 3 peaks of activity were recorded (figs 49 and 50). Some of the night activities were observed to be influenced by the presence of the brood (Figs 47 and 49 - 51), the activity pattern of the worker caste and to a lesser extent the soldiers. The findings on 25th December, 1976 indicated that as man} as 293 larvae and 192 pupae were found (table 13) and the corresponding activity pattern was marked (^ig 47) • On 1'8th February, 19779 as many as 415 larvae and 558 pupae were recorded from a nest (table 13) and this was found to have influenced the corresponding activity pattern in the nest under study (Fig 50). In this figure, many ants left the nest and many ants entered at any particular time which is higher than the normal activities found in the ants. Similar other recordings were made and are presented in figure 51* On certain days the ants were observed to be inactive all night. A close study of table 13 shows that there were only 9 larvate and 67 pupae which is by far very low as compared to the brood found 011 days that the ants were active. Figure $ shows that the ants were not as active as compared with nights such as in figure 47. The number of soldiers and workers recorded as foragers was few. - 85 - University of Ghana http://ugspace.ug.edu.gh Rainfall in the night seemed to have affected the activity of the ants hut the effect was not as narked as that found during the day (Fig 3?)* On a day 'when there v/as drizzling in the night (Fig 52), the anbs were active hut comparatively less active than the days without rain. On another day (fig 54) however, the rain started at 11 p.m and the activity of the ants dropped and the ants became active again when the rain stopped. On 30th April, 19779 t ^ percentages of larvae and pupae were high but the rains caused inactivity (fie 52). ■•3#JL Induced activity Observations listed above suggest that the activities of the ants were affected by the presence of brood, availa­ bility of food source, rainfall and the other factors such as temperature, light intensity and humidity* '*^ •5 Foraging space and distance The ants were found to travel over long distances in search for food and to forage on leaves. Recordings showed that the ants moved or covered different distances in the dry and wet seasons (table 15)* (a) get season During the wet season, the ants were found to travel longer distances to forage. On plot A, some ants went very far from their nests to forage and some even went into the other plot B to forage on the cultivated plants such as cassava and com. The others foraged within the savanna area — 86 — University of Ghana http://ugspace.ug.edu.gh IQ■POrH P< £o to 03 § MR O • CVJ 4 v o T~ c—a *r~ c— O r o r o • "d ” T— v o eh o o r t tQ 03 f t £3 r o r o r o r o r o ra WhO U i r \ O r o On o cvj£ -P • • • r o • • • •H Q) 00 x— VO • O v o c— to © a T“ ON ■%— IT \ CVJ CO r o t i o • t— ON ^4* c— v o v~ C§ D4 *1— CVJ r o CO T— o OJ ■*— ^aT orj c3 Eh -P 0 2 ^ -> o *H 02 3 q oi w B a> 2 -p ^ •H 05 -P rtf «H (U liLs. CQ <1) s*o u •3 -5 S 8 a.^ d o1 £«8v2. -P 03 ensis were already foraging. The ants found on - 87 - University of Ghana http://ugspace.ug.edu.gh plot A were found visiting plot B for foraging but those on plot B rarely went to plot A* Ants having their nests in areas with a lot of cassava plants were those that covered longer distances from their nests. The ants nesting in areas with com rarely moved more than 4 metres from their nests since most of the plants on which they foraged were found in the immediate vicinity, 6,3,4 Ants - -phytophagous insects relationship Certain types of sap-feeders were tended by CaywP,acvapimensis» These were Homoptera, aphids, larvae of stem boring beetles and coccids. The other association found were unimportant and occurred between the ants and scale insects (table 1o) , The ants were found to exhibit two types of association!: (a) Facultative associations - This type of association existed between the ants and the aphids, Aphis nerii Fonsc, ' The ants were found to collect honey dew from these insects. The aphids occurred on the bare surface of the leaves on which they fed. Tha ants usually spent only a few minutes around one aphid at a time. On several days, the aphids weafce left unattended by the ants. The ants exhibited similar relations with the scale insects that were found attached to stems of grasses. The time spent with the scale insects toas quite short, T7ith the Coccid, Pulvinarja sp. Boh,, found on cassava, the ants visited the coccids once a while and never spent more than 30 seconds with a coccid excepting in rare cases. - 88 - University of Ghana http://ugspace.ug.edu.gh With the Pseud©c®ccidae (Bysmicoccus breuipes (Clctti), Phenac®ccus madeirensis Green and Ferrisia virgata (Ck.lL) the ass©ciati©n was very passive. (b) Obligate associationship - This was only found on plantain and com and was so considered because the ants were beneath sheaths serving like tents to protect the sap-feeders that they tended. In some cases, the ants sealed the opening left with soil. Such an association existed between the ants and the larvae of the beetle Garpophilus fumatus Boh* which fed on the steins of com and also the aphid Pentalonia nifrronervosa Ooq. on plantain suckers• 6o3»5 Food items collected by ants The ants were found to be moving on almost all plants in their foraging space but were found to spend more time on certain species especially Zea mays (Com), Musa spp. (Plantain and Banana), Euphorbia tirucalli, Cocoyam and Solanum melongena. On certain plants they were only found when such plants had sap-feeders on them (table 16). On certain "plants such as Zea mays, their activities caused markings on the leaves as areas constantly gnawed became pale green and later whitish* On Euphorbia tirucalli, the ants were found on the floral parts and spent several minutes on them moving from one flower® to the next. Their presence on Musa spp. was mostly for tending aphids and Homoptera that were found o$ these plants. - 89 - University of Ghana http://ugspace.ug.edu.gh _ 9C - When the ants were sprayed and prevented from visiting the sap-feeders, a few were affected. Information on this is provided in table 16. On Solanum rnelon^ena, the ants collected floral parts mainly petals, pollen grains or anthers and sometimes excret*,- from the Tingid Urentius hvstricellus (Richter). 6.3.7 Food items These ants were observed to collect insects as food items into their nests. The greater number of insects collected were termites, mostly Microtermes spp., when the nests of such termites were destroyed by farmers or other environmental factors. Parts of other insects such as wings of butterflies, seeds of grasses, excreta from aphids and larvae of certain beetles formed the bulk of the food that the ants carried into their nest. On certain plants such as Euphorbia turicalli, the ants collected nectar. Examination of the gut contents of ants that spent hours on this plant revealed the presence of a solution which was similar to the secretions that the plant produced. Other food items that the ants were found collecting were the anthers of com and the floral parts of Solanum melongena. Parts of the stem of com and other plants are usually found gnawed by the ants but it is not known if these ?;ere used as food or for building of tents over sap-feeders. University of Ghana http://ugspace.ug.edu.gh Ta bl e 16 : Pl an ts on wh ic h an ts we re fo un d fo ra gi ng (d am ag e an d ma te ri al s co ll ec te d) *d 4 0 - p ctf o a 0 rH r -| £ o ro O 0 CQ 'c$ H ro cti •H 5p 0 0 0 aJ o (H -P •3ro H P h i> ctf a j 0 0 ►3 02 CQC d & Oo 03 •PO0 CQ .3 O a H O O M O CQ £ »§ s0£o r£ £ £:0 0 0 nd •Xl o3 S S to -P 0 0 0 o £ £0 o O o d ,3 Hr^i 0 O 0 O CQ CQ 0 r “Jo CQ0} r H 0 0 > r £ £ j aJ 02 M 0 *■3 *d00 fe- ICQffiao - P P=> CQ - P•H s £]o 0o•H "I 3 5 t3 3 dro r H 0 0ft £ •3 - P I I 0O • H •n CQ 02 - P - P • H • H r-» Cm ■ s n d .Cl CQ f-l 02 CQ 0 0 0J 4 > > O aJ CtJ d 0 03 CQ r H r H CQ 5 0 i—I 02 - P•r) g © 0 - P CQ CQ •H r r l f t ' £ C0 0 ro 0 n d 'd t o CQ !>> r H >3 • H 0 crt 0 » d P - p £ o 0 O 0 «—1 r-'—l P < - p • CQ - P•H I *3 s alH H rO P< 0 g <“ i r-> 60 0 d K ft O •H O • 1-3 X Q H a j h = i c\5 • • i— ! O d 0 1-3 Ca 4 9 • ctf • H P i O O £ H i Q » d CEJ 0 CQ 02 •H 0 cd 0 O . 0 ,g * H£J ■3 1 • H 02 • H • r l n d 0 - P • O -1-5 5 r H - P cd •H 02 a * d - p • H 0 c6 CQ 0 0 • H c 3 'S id s | CQ § r p a ro o3 m Ct» U O p . •3 0 3 O • H 0 OX , r3 CQ c3 •r-i § • HCQ g aS P I csj CQ aJ C ;' r H P O 0 M 1 CQ j g « r * 1 S j d0 5 Pa ss if lo ra fo et id a L* Pa ss io n fl ow er le av es an d fl ow er s no da ma ge Bo er rh aa vi a di ff us a L , le av es no da ma ge University of Ghana http://ugspace.ug.edu.gh Table 17: Monthly catches of alates of Camp*ac vapixaensio MONTH ‘■IfiLSS jt&iales TOTAL September, 197 ^ 14 6 20 October, 15 3 18 November 4 0 4 December 1 0 1 January, 1977 0 0 0 February 3 0 3 March 17 3 20 April 39 5 44 May 47 2 49 June 19 88 July 4: 0 4 August 3 1 4 September 17 4 21 October 13 7 20 University of Ghana http://ugspace.ug.edu.gh vo 6.3*8 Flight activity Light trap catches of alates indicated that the number of ants probably flying before 6 p.m were many and as such very few were caught between 6 p*m and 6 a*m (table 17). 6*4 DISCU3SI0I) *4.1 Fora^ in/? activity of Camp.acvapimensis ■The ants were active during the day a,s well as night. Daring the day they collect only food and at night the activity mostly dealt with food collection, nest cleaning and nest expansion. Foraging activities generally involved workers, occasionally soldiers were recruited to help in food collection when they were found to act as food carriers or guards during food collection* Similar findings were reported by Dobzansky (195S) in the genus Formica where the foragers recruited other foragers to the food source. In Camp.acvapimensis, all the castes have the characteristic raising of the gas ter and this possibly signals the detection of the food source to the other foragers. The diuma.l activities of the ants seemed to be affected by temperature and humidity. From the results, it is evident that these environmental factors influence the activities of the ants. Lower temperature and high humidities tend to influence the activities of the ants in that the ajits are more active under such conditions and - 91 - University of Ghana http://ugspace.ug.edu.gh are inactive when these factors vary. The only factor that tended to rule out the effects of these two factors was availability of food source in the environment. Availability of food during a very high temperature induced, the ants to continue foraging and collect food. The ants were active on such sunny days, collecting food into the nest tfhich was an indication that the presence of a food source in the environment influenced foraging activity of the ants. #hen food 7/as available in the environment, then humidity and temperature lost their effects on the ants. Another factor that influenced the activity of the .ants was presence of brood in the nest. These ants were active when the larvae found in the, nests were many. This agrees with the findings of Towles (1S52) and Schneirla (1944) > who reported that the presence of brood and the type of brood in the nests had effect on the activity of the ants. In the present finding, the larvae seemed to influence the activity and this is probably due to the inactivity of the pupae and the active feeding nature of the larvae. On the whole the ants were found to be active during the periods when there was occurrence of high percentages of brood (larvae)* Ackonor (1977) reported of Cat.yqxlneensls as maintaining a high level of foraging activity throughout the year. This he attributed to the occurrence of brood and alates in the nest throughout the year. University of Ghana http://ugspace.ug.edu.gh - 93 - I n Camp, acvaplrnensis» the presence of brood does not influence the diurnal activity of the ants but instead the night activities are affected. This is an indication that the percentage of brood in the nests influenced the activity of foragers up to a point. Another factor influencing the activity was the presence of food source in the environment. Teal 1 is (1963) associated the functions of the worker caste with hunger* In Gamp♦ acvapimensis the presence of food source influenced the ants. On days with high light intensities and very low humidities, the ants were seen foraging continuously because of a broken termite mound* Such foraging activity defined the humidity and temperature as factors influencing the activity of the ants. Ackonor (1977) also noted that the return of foragers which had been successful in obtaining food brings more stimulation to the inmates of the nests to urge them out in search of food* This is probably the cause of the numerous workers leaving the nes± of Camp * acvap imens is to forage for food and also probably the raising of the gaster also serves as a signal for the presence of a food source. Adabie (1977) attributed the stimulation of foragers in the nest of Crematester depressa (Latreille) to the returning of foragers and the probable scattering of the developing larvae in the lower chambers of the nest* She felt that all returning foragers University of Ghana http://ugspace.ug.edu.gh entered the lower chambers and stimulated more workers to forage and increase food supply. It is therefore not surprising for the first individuals of Gamp.acvai»imensis to return into the nest to raise their gasters as a signal causing more workers to leave the nest to the food source* On days that rainfall seemed to be oa influencing environmental factor, Camp. ac vapimensis remained active during the rain on leaves. Other ants retreated into their nests such as was found in Camp, sericeus (personal observation) and C a t«;?u in e en s i s in which Ackonor (1977) observed retreat during the rains. This is an indication that the effects of rain on the activity of Camp.aovapimen3is is negligible. Temperature and humidity were found to be influenced by changing seasons such as the harmattan. The ants were very inactive all day and those ants that cane out of their nests did not move far from the nest but immediately went back into their nests. This was rather unusual as the 1 ante forage far from the nest as a habit even in the dry season. Harmattan was therefore a factor that affected the general activity of the ants. 4-V-l-Z — 0tumal activi ty Effects of environmental factors on the nijht activi­ ties were not marked. Even when the humidities remained high and temperabures low but the ants were found to be active. - 94 - University of Ghana http://ugspace.ug.edu.gh There is the possibility that the ants have preference for high humidities and low temperatures* This is very likely because on days v/hen clouds caused low temperatures, the ants were seen active. The fact that these ants were avoiding sunlight and went into their nests when light intensity becam4 high is an indication that low temperatures favoured their activity. This possibly accounts for their increased activity on all nights. 6«4<>2 Foraging space and distance Foraging space and distance of Camp, acvapimensis was influenced by the seasons. During the wet seasons, the ants were found near the entrance of their nests. In the dry season, the ants travelled over long distances in search of green vegetation on which they foraged. Such movements might have been influenced by the type of food the ants were searching for. This is because the ants were alwa,ys searching for green vegetation. The subsequent scrappings of the stems of the green plants caused a concern other than a mere utilization of such materials for building tents over sap-feeders as these were not found during the dry season, and the ants were not building numerous tents over sap-feeders in the savanna areas as they do in other places. It is possible that the ants use such materials for food* Wyniger (1962) reported on Solenopsis geminata (F) as gnawing on plants and licking the sap of those plants. - 95 - University of Ghana http://ugspace.ug.edu.gh It is probable that this same behaviour is exhibited by Camp, acvapimensis which was reported by Evans (1973) as using such materials for the construction of tents over the Mealybugs which they tended. Sucking of plant sap seems to be a common behaviour of certain ants for Ackonor (1977) also reported that Cat, guineensis possibly was sucking sap from apical tissues of shoots and also collected materials from plants. It is therefore not surprising that Camp, acvapimensis also collects floral parts from certain plants and sap from others as source of their liquid food. Tending of other insects for honey dew is probably to supplement their supply of liquid diet. It seems that more is involved than a mere nest building habit reported by EVans (1973)* In other ants such as Crem, depressa, Adabie (1977) reported that the wood fibres used for nest materials were scrapped from the backs of cocoa pods and other parts of the tree. A further inves­ tigation into the use of sap and other plant materials by Camp, acvapimensis will throw light on the dama.ge that the ants cause. In the savanna areas, such plants were not available to stimulate further investigations. This may account for the constant search for green plants by Camp, acvapimensis. The distance travelled by these ants in the dry season therefore was in search of green plants to obtain sources for their food rather than wandering aimlessly. - 96 - University of Ghana http://ugspace.ug.edu.gh vo Since these plants weote farther away, the ants were found several metres from their nests* Another possibility was that the sap-feeders which they tended were on green plants and such plants were far’ from their nests and they had to cover all the area in search of these sapr^ee<3-ers* The occurrence of sedondayy nests in areas away from the primary nests was also a possible reason for the distances that the ants moved* *4*3 Pood items collected by Camp*acvapimensis The main food items collected by these ants were termites and other insects* This collection of insects confirm the report of 17h.eeler (1922) who stated that the ants of the genus Camponotua collected unicellular insects and the excreta of aphids for food* It is therefore not surprising for Camp * acvap imensi s, a member of this family, to collect termites for food and tend aphids and other insects in the savanna areas. The food of this ant included honey dew collected from the aphids and coccids that it tended* Their continued visits to the aphids is an indica­ tion that the aphids secreted the honey dew in considerable quantities and the ants made use of them* Plant Juice seemed an important component of the diet of these ants. This can be inferred from the constant sucking of the juice of the seeds of Neem trees and mangoes* The other juice source was nectar from Shphorbia tiri.icalli* Certainly, these ants - 97 - University of Ghana http://ugspace.ug.edu.gh - 98 - might look for nectar from other sources when plants that give off such nectar are not available* The gnawing of pods, as has been pointed out earlier by Evans (1971)> may be for a different purpose in the cocoa farm as Mabie (1977) pointed out in Crem.depressa that the ants use such materials for nest building and Ackonor (1977) also found that Oat. guineensis were sucking sap from the apical tissues of the shoots as a source of liquid food. In the savanna, this was probably serving in different purpose other than tent building. The report of Evans (1973) only attributed this behaviour to the nest building habit since they were concerned with finding the source of the pd‘d rot disease. This was most likely forming part of the diet of the ants because Gorenz (19S9) reported that Gamp.aovapm ensis gnaw the skin of cocoa pods for nest building. The obvious reason for these ants to use the gnawed materials for food is that in the savanna areas very few tents are constructed over aphids and other insects in the dry season. Secondly, the primary nests of these ants were in the soil and the use of sap and gnawed materials for incorporating in the nests was not found. However, in the dry season the ants constantly visited green plants and removed portions of the stem. Such materials could be used for food rather than nest building. Coupled with the fact that these ants suck juice from fruits and nectar from flowers, it is University of Ghana http://ugspace.ug.edu.gh possible that these ants were looking for such sap as a food requirement, Wyniger (1?62) reported on the gnawing behaviour of Solenopsis Deminata and how it licked the sap that came out* It is very likely that Camp*aovapimensis was also licking sap from the plants from which they gnawed the materials. Seeds collected from grassed were transported and stored in one or more cells in the nest. The food items commonly found in excavated nests were seeds, anthers and pieces of stick. No termites or insects were found in the nests. This is probably because the termites and other insects were used immediately they were carried into the nests. The probable reason is that these insects could not be stored for longer periods without decay but the seeds could stand storage for several days. This accounted for the absence of such food items from the nests. 6.4*4 Ants phytophagous insects relationship The association between these ants .and the sap-feeders were facultative. The aphids and coccids were tended without the ants building tents to protect them but when these were tended under th€ sheaths of com or plantain, the sheaths protected the sap-feeders. The only associa- tionship which could be called obligate was that between the ants and the larvae of the beetle in which the ants constructed tents to seal off the openings, left. - 99 - University of Ghana http://ugspace.ug.edu.gh In this, the ants used soil to seal the space between the sheath and the stem of the plant. During pupation of the beetle, the ants left the sheaths* In some cases, some aphids were found in the same sheaths and the ants continues to tend such aphids. The association found between the aphids and the ants depended on where the aphids were found for when they were found on leaves as they occured on cucumbers, the ants tended them and did not afford them any protection. The damage caused to most of the plants on which the ants were found tending the sap-feeders was jointly caused by the ants and the sap-feeders. 6.^.5 Flight activity The absence of alates from light trap catches after 7 p.m. was a possible indication that the alates were probably flying off at an earlier time. The number of alates in the nests were also very few and this could mean that colony founding might involve colony expansion rather than increase by the number ol alates• This is accounted for by the always high percentage of larvae and pupae of the worker and soldier castes as compared with the alates that were found in the various nests that were excavated. - 1 0 0 - University of Ghana http://ugspace.ug.edu.gh Generally, flight activity was influenced by the onset of the rains as the ants were seen in light trap catches some days after the rains* This is probably due to the size of the alate queens as they were found to be larger and needed time to come out of such nests* The male alates were seen in the light traps early probably because of their small sizes and this probably causes their easy movement out of the nest than the female alates. University of Ghana http://ugspace.ug.edu.gh - 102- S T J M M A H Y 1, The study deal with the distribution, nesting habits, preferences, seasonal population pattern, general activity patterns such as foraging and food collection and economic importance of Camponotus acvapimensis in savanna areas of Accra, Ghana* 2* The factors which contribute to the distribution of Camp»flcvapimensis were investigated by observations, surveys and field experiments. The distribution, abundance, activities and nesting habits were studied on 2 plots, one plot was a farm land and the other a grassland each measuring 1 hectare* The two habitats afforded basis for comparison within a typical savanna ecosystem. 3* In savanna grasslands, the ants were found in areas with poor shade provided by climbers and a few shade plants, mostly annuals (grasses)* The ants were almost absent in areas with continuous shade. Occasionally, their populations increased in areas without shade when there was drought. Light intensity, availability of food and other climatic factors ©ere also found to influence distribution* 4* Nest distribution assessment made in 4 different areas of no-shade, broken shade, poor shade and continuous shade showed varied statistically significant differences 0 : H A P T S B 7 University of Ghana http://ugspace.ug.edu.gh - 103 - ■between such shade regimes, 'The ants were found to forage in any of such areas even though their nests were numerous in poorly shaded areas. Further investigation revealed that the ants abandoned the poorly shaded areas when the poor shade condition was lost. It was concluded that Camp*“vcvapimensis loves poorly shaded arefes but may live in broken shade and no-shade areas as well depending on changing seasons. 5. The colony of the ants was not extensive and was made of one primary nest (usually in the soil) and a few secondary nests (in which the ants tended sap-feeders). Such nests were near the primary nests but usually depended on the distribution of the vegetation. 6. Most nests were found in the soil, grass roots and trees and those in the trees were few. Distribution with respect to shade regimes was influenced by the seasons and the nests were numerous in no-shade areas after bush fires, Nests were mostly found in soils with excess of clay and silt but were rare in sandy soils. 7o Fights between soldiers of different colonies were short-lived and involved few soldiers dashing at members from other colonies, giving quick bites with their mandibles and retreating quickly. Only about T0?S of the soldiers we£e engaged in guarding and fights. This represented only about 2/J of the population. ITo harm University of Ghana http://ugspace.ug.edu.gh 8* Camr«apvapimensis was found to occur to g e th e r w ith o th e r a n ts . They c o -ex is te d w ith Camp^g e ric eu s b u t excluded them from fo rag ing a re a s . However, th e two a n ts were found a t tim es fo rag ing on th e sarae p lan t* In some n e s tin g a re a s , Cami,*acvapimensis was found n e s t in g in the s o i l near Annoma d r iv e r a n ts . Gamp■ acvapimensis workers appeared d is tu rb ed by P la ty th y reu s ta rs a tu s 'flhich ac ted as i t s p red a to r . 9 0 The an ts were found to excavate n e s ts in th e s o i l o r g ra ss ro o ts and th ese had numerous c e l l s . The number o f c e l l s ranged from 3 - 1 2 and t h e i r v e r t ic a l depth ranged from 12.3 cm - 63.1 cm. The tu nn e ls were m ostly in c lin e d a t 60° to th e h o r iz o n ta l s u rfa c e . The c e l l s were la rg e r n e a re r the main en trance o f th e n e s t bu t narrowed deeper down. The la rv ae were in c e l l s deep do mi and th e pupae n e a re r the su rfa c e . The number o f c e l l s had a b ea rin g on th e con ten ts (brood and c a s te ) o f th e n e s t . Secondary n e s ts were w ithout brood though a few had 1 0 - 3 0 pupae on ly and in such n e s ts th e an ts tended th e sa p -fe ed e rs . Some secondary n e s ts found under c reep ing g ra sse s o r heaps o f decaying le av es had on ly pupae and in r a r e cases few larvae* Some secondary n e s ts had male and female a la te s a s w e ll. P rim ary n e s ts had la rv a e , pupae, workers, s o ld ie r s and winged forms ( la rg e r femal.es and sm alle r males)* was done to any of the ants involved. University of Ghana http://ugspace.ug.edu.gh 10* The population of ants had the highest peak of 1,572 individuals in February and a minor peak of 916 in June* The highest peaks were both recorded in periods without rain and the lowest peaks in periods with rain. The ants abandoned’ their nests when the nests r/ere partly excavated during digging by farmers or accidental excavation of farm habitats* Ttihen refuse collects at the entrance thus blocking the entrance, the ants also abandon such nests. 11* The ant was found to be active both day and night but /— - —— - activity in the night was marked as compared with that of day* Several peaks of activity were observed in the night* Nest cleaning took place during the night* Day activities were influenced by light intensity, temperature and humidity but light intensity and temperature determined the pattern of activity. Morning activity was around 6*20 - 10 a*m and afternoon activities started around 3 — 5 p#m* Days with high light intensities had low activities and days with low intensities had increased activity. Humidity was average* Rainfall had no effect on activity pattern but activity increased considerably after each j rainfall. Availability of food sources increased the activity of the ants and was the only factor that made the ^ ants defy the effects of temperature and light intensity* Effects of presence of brood on activity were marked during the night activities but did not influence the day-time activities* - 105 - University of Ghana http://ugspace.ug.edu.gh 10* The population of ants had the highest peak of 1,572 individuals in February and a minor peak of 91 o in June. The highest peaks were both recorded in periods without rain and the lowest peaks in periods with rain. The ants abandoned their nests when the nests were partly excavated during digging by farmers or accidental excavation of farm habitats* Tihen refuse collects at the entrance thus blocking the entrance, the ants also abandon such nests. 11* The ant was found to be active both day and night but ------------ activity in the night was marked as compared with that of day* Several peaks of activity were observed in the night* Nest cleaning took place during the night. Day activities were influenced by light intensity, temperature and humidity but light intensity and temperature determined the pattern of activity. Homing activity was around 6*20 - 10 a.m and afternoon activities started around 3 - 5 P*m« Days with high light intensities had low activities and days with low intensities had increased, activity. Humidity was average. Rainfall had no effect_on activity pattern but activity increased considerably after each rainfall. Availability of food sources increased the activity of the ants and was the only factor that made the ( j ants defy the effects of temperature and light intensity* Effects of presence of brood on activity were marked during the night activities but did not influence the day-time activities. - 105 - University of Ghana http://ugspace.ug.edu.gh 12. The ants foraged long distances from their nests singly and their foraging space covered an area of about 2137.2 sq. metres. Foraging was done on vegetation and the ants climbed plants purposely for tending coccids and other sap-feeders or collecting juice from the fruits. Food items carried into nests included termites and seeds of grasses. Sugary materials were carried in liquid form. The ants caused injury to the flowers and leaves of Solanum melongena stems of com and some other cultivated plants. 13. The ants were found tending the aphids Aphis nerii, passively and Pentalonia nigronervosa activelya Other sap-feeders tended are the coccids Pulvinaria sp., the Pseudococcidae Sysmicoccus breuipes, Phenacoccus madeirensis and Ferrisia virgata. Other important partners were the larvae of the beetles Carpophilus f umatus. 14* Flight activity occurred about four or five days after the rains and the ants were mostly found during the early hours of the night but were rarely seen in the late hours of the night and before dawn. The male alates were seen some days before the females appeared. The number of male alates was always more than the female alates. University of Ghana http://ugspace.ug.edu.gh R E F E R E N C E S University of Ghana http://ugspace.ug.edu.gh R E F E R E N C E S Ackonor, J.B. (1977) The distribution, nesting habits and activity pattern of Cataulaucus guineensis P. Smith (Hymenoptera! Formicidae) in a Ghanaian cocoa farm, M. Sc. thesis University oft Ghana Legon p 137“1 44* Adabie, D.A. (1977) The distribution, activity and nesting habits of Crematogaster depressa (Latreille) (Hymenopteral Formicidae) in a Ghanaian cocoa farm, M.Sc. thesis University of Ghana Legon 154PP* Anipare, C. (1971) B.Sc* project, Dept, of Zoology University of Ghana Legon. Axyeetey, E. A. (1 971) The biology and ecology of Macromischoides aculeatus aculeatus (Hymenoptera! Formicidae) in a cocoa farm in Ghana* H* Sc. thesis University of Ghana Legon 185p Ayre, G. L. (1958) Some meteorological factors affecting* the foraging of Formica subnitens Creighton (HymenopteraJ Formicidae)* Insectes 3oc, 51 2 I 1 47 + 157 Bigger, M. (1972) Varietal susceptibility to attack by cocoa insects. Proc. of 4th Qonf. of West African Cocoa Eat* 1 974 PP Bolton, B* (1 973) The ant genera of West Africa* A synonimic synopsis with keys (HymenopteralFormicidae) Bull* Mus* Nat. His. London 27 (6), 317 “ 368. Brian, M.V* and Downing, B.M. (1958) Nest structure of some British ants* Proc* Xth Int* Cong* of Sht* Montreal 2, 539 - 558* - 107 - University of Ghana http://ugspace.ug.edu.gh - 108 Brown, E. S. (1959) Immature nutfall of coconuts in the Solomons Island. I - Distribution of nutfall in relation to the ant Amblypelta and certain species of ants* Bull Ent Hes. 50* 97“133>523~558« De Geer (1884) Camponotus pennysylvanicus permeated by a fungus* Leidy* Philad* 1884> P*9* Dennis, C. A. (1938) The distribution of ant species in Tennessee with reference to ecological factors, Ann* Sht* Soc. Amer. 31 (2) : 267 - 308. Dobzansky, J (1 958) Partition of foraging grounds and modes of conveying information among ants. Acta. Biol. Exp. 181 55 - 67j cited by Wallis (1 964) Dogel, V. A. (1924) Qualitative analysis of the fauna of meadows in Peterhof Researches in the quantitative analysis of terrestrial fauna. Rev. Zool. Russe 4 0 - 2), 117 - 154. Dutta, A* C. (1 964) Botany for Degree students Oxford Univ. Press p 238 - 241. Emery, Ce (1877) Saggio di un Ordinamento naturale dei Mirmicidei, conaiderazioni sulla filogenesi delle Formiche. Bull, ait. Ital. 99 PP 67 - 83 pi 1• finery, C. (1883} Camponotus sylvaticus. Ann. Mus. Genov. 18 pp 448 - 452* finery, C. (1896) Saggio di catalogo sistematico dei generi Camponotus, Polyrhachis affini. Mem. Accad. Sc. Bologna 5 I 761 - 780. University of Ghana http://ugspace.ug.edu.gh finery, C. (1899) Camponotus aqwapimensis. Ann. Mus Civ Genova 39, P 501 fineiy, C. (1 901) Camponotus akwapimensis.Bull. Soc. Eht. Italiana. 33, P 63* finery, C. (1920) Le genre Camponotus Mayr Nouvel essai de sa subdivision en sous-genres. Rev* Zool. Afric. Bruxelles. 81 2 (1920) pp (229 - 260). Entwistle, P. P. (1972) Pests of cocoa. Tropical Science series Longman group Ltd. 779PP Evans, H.H. (1971) Transmission of pod rot of cocoa by invertebrates. Nature, London. 2321 34-6 - 347* Evans, H.C. (1975) Invertebrate vectors of Phytopthora palmivora causing black pod disease of cocoa in Ghana. Ann. Applied Biology, 75 • 331 - 345* Firerapongj S. K. (1 975> ^be biology of the black citrus Aphid Toxoptera aurantii Boy on cocoa in Ghana, M. Sc. thesis, University of Cjhana, Legon, Forel, C. R. ('1890) Camponotus species of rubripes and varieties noticed, Ent, Belgo. 18 $6 Forel, C.R. (1912) Formicides noetropiques Part VI sous famille camponotinae. Mem. Soc, Ent. Belgique, 201 59-92. Porel, C, R. (1914) le genre Camponotus Mayr et les genres voisins Rev. Suisie Zool. 221 257 - 276 Goetsch, W. (1929) Seed collection. Naturwiss 17 I 221 - 226. - 109 - University of Ghana http://ugspace.ug.edu.gh Gorenz, A Gosswald, Las ton, D, Leston, D, Leston, D, Majer, J. Majer, J. Majer, J. Majer, J. M. (1969) Spread of Phytopthora pod rot from the tree base to pods in the canopy. Ann. Rep. Cocoa Res. Inst. Nigeria 1 968 - 1969 * 53 - 54* K. (1932) Oekologische studien uber die toeisenfauna des mittleren Mainbiestes . Zeitsch. Wiss. 2oll. 142 (1) : 1 - 56. , and Hughes, B. (1968) The snakes at Tafo, at forest cocoa farm locality in Ghana. Bull, de I.I.F.A.N. 50 (A2) J 738 - 770. , (1 968) Diurnal activity of Oecophylla longinoda. Rep. Cocoa Res. Inst. Tafo. 1965 - 19661 62 - 72. > (1971) Ants, ca,psids and swollen shoot in Ghana, interactions and implications for pest control. Proc. 5rd Int. Cocoa Res. Conf. Accra. 1969 205 - 221. D. (1972) The ant mosaic in Ghanaian cocoa farms. Bull. Ent. Res. 62 5 151 — 160. D. (1973) The nature and role of the ant complex in Ghana cocoa farms. Pfr. D. thesis Univ. of Ghana Legon 553pp. (1974 ) ‘^ ie use ant in an intergrated control scheme for cocoa. Proc. 4th Conf. of West African Sit. 9th - 13th Dec. 19741 181 - 190. b (1976a) The ant mosaic in Ghana cocoa farms, further structural considerations. J. Appl. Bcol. 131145-175. University of Ghana http://ugspace.ug.edu.gh Majer, J. D, (1 976b) The influence of ants and ant manipulation on the cocoa farm fauna# J» Appl• Ecol, 15* 145“*155* Majer, J* D. (1976c) The maintenance of the ant mosaic in Ghana cocoa farms, J, Appl, Ecol. 13 I 123 - 144* Mayr, C, (1862) The genus Camponotus acvapimensis Verh, Zool, Bot, Ges,Wien,12 p 664 Oertzen, V, H, (1887) Appendices a nom memo ire sur les sensations des insectes, Bo E, Z* 32 • 255 - 26$* Reh, L, (1905) Camponotus akvapimensis Zeitsch, f, Pflanzenkrankh, 15> P 134. Room* P, M, (1971) The relative distribution of ant species in Ghana !s cocoa farms. J, Anim0 Ecol* 40* 5^9 "535* Room, P* M* (1974) Arthropods taken on the mistletoe Tapinanthus bangwensis (Engle and K* Krause) growing on cocoa in Ghana* J, Anim. Ecol. Schneirla, T* C. (1 944) The reproductive fluctuations of the army ant queen as pace-maker of the group behaviour pattern* J, New York Ent, Soc, 521 153 - 192. Smith, P. (1866) Notes on Hymenoptera, £ht* Ann* for 1866 pp 122-137* Sudd, H* J. (1967) -An introduction to the behaviour of ants* Edward Arbold(publishers) Ltd, London, 50 - 135* Tabolt, M* (1 934) Distribution of ants in the Chicago region with reference to ecological factors and physiological toleration, Ecol0 35 • 416 - 439* Tabolt, M* (1943) Population study of the ant Prenolepis imparis Say. Ecol. 24 I 31 — 44* - 111 - University of Ghana http://ugspace.ug.edu.gh Vbwlesj D, Wallis, D, Wheeler, Wheeler, Wray, D* Wyniger, R » M# (1955) The foraging of ants. Br# J# Anim# Beh# ( i# (1 965) The relation between worker function in ant colony 0 Proc# Zool# Soc# Lond# 139 I 589~6$5* ro M# (1 922) Ants of the Belgian Congo Expedition# Bull# Ame# Mus. Nat# Hist, 45 J 187 - 190# ro M# (1930 Prenolepis imparis Say. Ann# Ent# Soc# Amer# 23 I 1 - 26. .. (1938) Notes on the Southern harvester ant (Pogonomyrmex badius Latr.) in North Carolina Ann. Eht# Soc# Amer# Columbus 31 I 196-201 # # (1962) Pests of crops in warm climates and their control# Basel Verlag fur Recht und Gesellschaft (Acta trop# supp# 7 ) 4^5 -475 • University of Ghana http://ugspace.ug.edu.gh A P P E B D I X T A B L E S OiSE - S I X T Y University of Ghana http://ugspace.ug.edu.gh A P P E N D I X T A B L E S 1 - 60 Table Meteorological data* at study Area 1 Head measurements of some Soldiers 2 - 7 Head measurements of some Workers ** 8 - 1 3 Diurnal activity patterns . 0 1 4 - 3 9 Nocturnal activity patterns * I . * 40 - 60 University of Ghana http://ugspace.ug.edu.gh Table 1. Meteorological information on the study plots during the study period. Month Amount of rain(mm) No. of rainy days July 23.4 5 August 0.8 1 September 33.6 4 October 81.3 8 November 30.0 3 December 8.9 2 January 15.2 1 February 11 . 0 4 March 55.1 2 April 86.4 9 May 85.7 9 June 71.7 15 July 9. 6 8 August 18.2 7 September 36.4 --------------------------------------------------- -— ------ ------------ 6 University of Ghana http://ugspace.ug.edu.gh Table 2 j Head measurements of soldiers in certain nests of C. AQvapimensis m HL m HL 1.81 2.23 1.89 2.31 1.77 2.19 2.23 2.50 1.81 2.31 1.89 2.27 1.62 2.27 2.73 2.42 1.81 2.23 1.85 2.27 1.75 2.27 1.89 2.12 1.85 2.31 1.77 2.27 ■1.96 2.39 1.67 2.19 2.04 2.42 1.87 2.39 1.85 2.42 1.89 2.42 1.92 2.42 1.89 2.42 1.92 2.15 1.81 2.19 1.41 1.85 2.U4 2.42 1.74 2.19 1.89 2.42 1.65 2.15 1.85 2.31 1.69 2.23 1.69 2.19 1.69 2.23 1.85 2.23 1.73 2.12 1.65 2.19 1.85 2.31 1.81 2.31 1.89 2.39 1.81 2.31 1.89 2.19 1.85 2.15 1.81 2.31 1.85 2.23 1.92 2.15 1.85 2.27 1.89 2.42 1.92 2.31 1.92 2.35 1.96 2.31 1.69 2.u8 1.75 2.19 1.92 2.15 1.77 2.19 2.04 2.31 100.09 124*70 University of Ghana http://ugspace.ug.edu.gh Table 3j Head measurements of C. acvatiimensis so ld iers from ce rta in nests of Nest 7 30/ 12/76 m HL m HL 1.69 2.00 1.81 2.42 1.75 2.25 1.89 2.50 1.69 2.13 1.69 2.19 1.69 2.25 1.63 2.25 1.50 1.89 1.89 2.42 1.81 2.50 1.69 2.42 1.63 2.13 1.69 2.42 i.50 1.89 1.65 2.35 1.63 2.19 1.81 2.35 1.69 2.25 1.69 2.23 1.69 2.13 1.75 2.08 1.81 2.25 1.75 2.19 1.81 2.20 1.89 2.19 1.89 2.19 1.75 2.50 1.75 2.06 1.75 2.42 1.75 2.00 1.89 2.50 1.89 2.50 1.81 2.19 1.63 2.00 1.81 2.31 1.50 2.06 1.63 2.15 1.6J 2.19 1.75 2.31 1.89 2.06 1.75 2.08 1.50 2.13 1.89 2.13 1.69 2.13 1.75 2.23 1.8i 2.75 1.69 2.23 1.75 2.25 1.63 2.25 1.89 2.19 1.89 2.50 1.85 2.42 1.96 2.42 1.92 2.42 1.42 2.15 1.92 2.19 1.75 2. 19 1.69 2.42 1.65 2.50 1.69 2. 19 1.69 2.42 1.85 2.50 t.65 2.(9 1.69 2.42 -.1*7? _ 2.23 University of Ghana http://ugspace.ug.edu.gh Table 4 : Head measurements of soldiers from certain nests of n. Hnvapimensig. Nest 8 11 28/2/77 OT EL HW HL 1.62 2.07 1.65 2.31 1.53 2.04 1.73 2.15 1.69 2.12 1.50 1.96 1.65 2.27 1.62 2.04 1.73 2.27 1.52 2.15 1.57 2.08 1.54 2.12 1.57 1.92 1.65 2.15 1.57 2.08 1.65 2.12 1.69 2.15 1.77 2.31 1.62 1.96 1.81 2.31 1.65 2.31 1.62 2.15 1.46 2.23 1.62 2.15 1.69 2.15 1.62 2.15 1.65 2.15 1.65 2.08 1.57 2.08 1.65 2.15 1.62 2.15 1.69 2.12 1.57 2.27 1.50 2.23 1.39 1.12 1.73 2.23 1.57 1.96 1.73 2.08 1.53 2.04 1.65 2.20 1.77 2.31 1.57 2.31 1.S7 2.04 1.73 2.04 1.53 1.96 1.81 2.04 1.73 1.96 1.65 2.20 1.69 2.31 1.77 2.23 1.77 2.31 1.65 2.08 1.73 2.12 1.53 2.12 1.73 2.08 1.73 2.31 1.65 1.96 1.65 2.27 1.62 2.07 1.65 2.04 1.69 2.23 99.36 University of Ghana http://ugspace.ug.edu.gh T a b l e 5: Head m e a s u r em e n t s of soldiers from certain nests of C. ^nimensis Nest 16 15/5/77 m HL m HL 1.81 2.23 1.85 2.31 1.77 2.19 2.23 2.50 1.81 2.31 1.39 2.27 1.62 2.27 2.73 2.42 1.81 2.23 1.85 2.27 1.73 2.27 1.89 2.12 1.85 2.31 1.77 2.27 1.96 2.39 1.69 2.19 2.04 2.42 1.85 2.39 1.85 2.42 1.89 2.42 1.92 2.42 1.89 2.42 1.92 2.15 1.81 2.19 1.42 1.85 2.04 2.42 1.75 2.19 1.89 2.42 1.65 2.15 1.85 2.31 1.69 2.23 1.69 2.19 1.69 2.23 1.85 2.23 1.73 2.12 1.65 2.19 1.85 2.31 1.81 2.31 1,89 2.39 1.81 2.31 1.89 2.19 1.85 2.15 1.81 2.31 1.85 2.23 1.92 2.15 1.05 2.27 1.89 2.42 1.92 2.31 1.92 2.35 1.96 2.31 1.69 2.08 1.73 2.19 1,92 2.15 1.77 2.19. 2.04 2.31 103.29 120 . . : University of Ghana http://ugspace.ug.edu.gh Table 6 s Head m e a s u r em e n t s of soldiers from certain nests of C . «™«T>imaEsis N e s t HW HL M HI 1.65 2.31 1.62 2.07 1.53 2.04 1.73 2.07 1.69 S.07 1.62 2.04 1.69 2.15 1.77 2.31 1.73 2.27 1.57 2.15 1.53 2.04 1.73 2.15 1,57 2.08 1.53 2.15 1.57 2.08 1.65 ZH5 1.69 2.15 1.62 2.04 1.65 2.23 1.77 2.12 1.39 1.96 1.73 2.23 1.57 2.31 1.57 2.31 1.77 2.04 1.65 2.20 1.69 2.04 1.96 2.20 1.57 1.96 1.73 2.23 1.57 2.08 1.65 2.15 1.73 2.08 1.57 2.27 1.62 2.15 1.69 2.12 1.39 1.12 1.73 2.23 1.53 2.04 1.73 2.04 1.69 2.31 1.57 2.08 1.77 2.31 1.65 2.04 1.73 2.08 1.53 2.31 1.73 2*12 1.53 2.12 1.65 1.96 1.65 2.27 1.62 2.07 1.65 2.08 1.53 2.04 145 2.20 1.69 2.04 1.65 2.20 1.53 1.96 1.73 2.23 1.77 2.31 1.77 2.23 1.57 2.08 1.73 2.20 101.54 i2 i .43 University of Ghana http://ugspace.ug.edu.gh Table 7 : Head measurements of soldiers from certain nests of C. acvftpimenais Nest HW HL HW HL 1.81 2.23 1.85 2.31 1.81 2.31 1.89 2.27 1.62 2.27 2.73 2.42 1.77 2.19 2.23 2,19 1.96 2.39 1.69 2.19 1.73 2.12 1.89 2.27 1.86 2.31 1.77 2.27 1.96 2.39 1.69 2.19 1.85 2.42 2.04 2.39 1.92 2.42 1.89 2.42 1.85 2.31 1.89 2.42 1.69 2.23 1.69 2.19 1.73 2.12 1.65 2.19 1.69 2.23 1.69 2.19 1.81 2.31 1.81 2.31 1.92 2.35 1.96 2.31 1.89 2.19 1.85 2.15 1.81 2.31 1.85 2.23 1.89 2.19 1.92 2.31 1.92 2.15 1.81 2.19 1.73 2J15 1.85 2.19 1.73 2.19 2.04 2.19 1.69 1.85 2.04 2.42 1.85 2.15 1.92 2.31 1.8 ° 2.42 1.92 2.31 1.92 2.35 1.96 2.31 1.69 2.08 1.73 2.19 2.04 2.31 1.77 2.19 1.92 2.12 1.65 2.23 1.42 1.85 2.04 2.42 1.73 2.19 1.89 2.19 113 .2 4 142.47 University of Ghana http://ugspace.ug.edu.gh Table 8. Head measurements of workers in various colonies of C. acvapimensis Nest 2 I5/1 0 /7 6 w H L H W H I 0.96 1.42 1 .22 1 . 3 9 0.96 1.46 0 . 9 6 1 . 5 4 1.04 1.42 0 . 9 6 1 . 5 4 1.04 1 .42 0.96 1 . 3 9 0.89 1.35 1.00 1 .27 0.96 1.62 0.92 1 .39 0.92 1 .46 0.92 1 .42 0 .89 1.35 1 .23 1.65 0.89 1 .23 0 . 8 9 1 .42 ’l.OO 1.27 1 . 0 4 1.39 1.00 1 .30 O .92 1 .27 1.12 1 . 5 0 O .92 1.30 0.96 1 .39 6 . 8 9 1.42 1.25 1.46 0 .96 1 . 5 4 0.96 1.58 0 . 8 9 1 . 5 4 0.96 1 .58 0 . 8 9 1 .46 1.00 1.42 0.96 1 . 5 4 0.92 1.42 1.04 1.23 0.85 1.42 0.85 1.46 1 .00 1.35 0.85 1.35 0.92 1 .39 1 .12 1 .27 0.81 1.27 1 .00 1 . 5 4 0.85 1.27 1 .00 1 . 5 4 0.85 1.30 0.92 1.46 0.92 1.42 1.04 1.39 0.96 1 . 5 4 0.96 1.58 0 .92 1.54 0.85 1.35 1.00 1 .30 0 . 9 6 1.35 52.07 78.77 University of Ghana http://ugspace.ug.edu.gh Ta b l e 9* H e a d measurements 0. acvapimensi6 o f w o r k e r s in N e s t 6 v a r i o u s c o l o n i e s 15/ 12/76 HW H L E W HL 0.96 1.46 1 .04 1 . 4 4 0.96 1 .54 1 .00 1.63 1.04 1.27 0 . 8 9 1.31 0.96 1 .39 1.13 1.31 0.89 1.27 0 . 8 9 1.25 OO• 1.35 0 .96 1.38 0.92 1.42 0 . 9 6 1.38 0.96 1.46 1 .00 1 .38 1.00 1 .39 1 .00 1 .62 1.04 1.27 1 .12 1.62 1.04 1.27 1 .04 1 .27 0.89 1.27 1.96 1.35 0.96 1 .35 0.96 1 . 5 4 0.92 1.42 O .96 1 .42 1.00 1 .54 O .56 1.42 0.89 1 .54 0.96 1.46 1.00 1 .39 1 . 0 4 1.42 0 . 9 4 1 .13 1 .06 1.25 1.00 1.56 1 .13 1,56 0.94 1.31 1 .04 1.25 0.89 1.44 0 . 9 4 1.31 1.06 1.44 0 . 9 4 1.31 0.89 1.25 0.96 1.31 0.89 1.25 0.96 1.31 0.96 1.31 1 .00 1.63 1.04 1.44 1.00 1 . 4 4 1.00 1.63 1 .04 1.63 0 . 8 9 1.31 1.13 1.31 1.06 1.44 0 . 8 9 1.25 O .96 1.38 0 .96 1.38 O .96 1.38 1.00 1.38 61.49 125.32 University of Ghana http://ugspace.ug.edu.gh Table 10o Head measurements of workers in various colonies of C* acvapimensis Nest 7 3Q/l 2/76 H W H L 1 . 1 9 1.62 0.92 1.46 0.92 1.42 1.04 1.42 1.00 1 .30 1.08 1 . 5 4 0.92 1 .27 0.96 1.46 0.96 1 .54 1.04 1 .50 0.85 1 . 3 9 1.15 1.58 0.96 1.46 1 .12 1.40 1.00 1 .30 0.96 1.35 0 .89 1.42 1.00 1 .50 0.92 1.27 0.96 1.42 0 .89 1.46 1 .00 1.35 1 .12 1.58 1.15 1.58 0.92 1.30 1 .00 1.42 1 . 0 4 1 .39 1 .12 1.58 1 .00 1 .30 HV7 H L 6 . 8 9 1 .27 O .92 1 .46 1 .00 1.30 1 .04 1 . 3 9 0 .92 1 .23 1 .12 1.46 0 . 8 9 1.46 0 . 9 2 1.46 0 . 9 2 1 . 5 4 1 .00 1 . 5 4 0 .92 1 .39 0 . 9 2 1.39 0.85 1 .30 1.00 1 .30 0 . 9 2 1.30 0.96 1 .39 1 .00 1 . 3 9 0.96 1.46 0.81 1.23 1 .00 1 . 5 4 0.92 1 . 3 9 1 .00 1.58 1.08 1 . 5 4 1 .12 1 .54 0.96 1.42 1 . 6 0 1 .30 1.15 1.58 1 .15 1 . 5 4 1 .12 1.54 57.54 82*23 University of Ghana http://ugspace.ug.edu.gh Table 11. Head Measurements of workers in various colonies of C. acvapimensis ITest 11 28/2/77 W HL TC E i l H L 0.96 1 .42 0.85 1 .27 1.12 1.39 1 .00 1 . 5 4 0.96 1.39 0.85 1 .54 1.12 1 .42 0.85 1 . 5 4 0.96 1.46 0.85 1 .30 1.04 1 . 5 4 0 .92 1 .46 0.96 1.54 0 . 9 2 1 .42 1.04 1.42 1 .04 1 . 3 9 0.96 1.54 0 . 9 8 1 . 5 4 0.89 1.25 0.98 1 .58 1.00 1.62 0.85 1.35 0.92 1 .39 O .92 1.46 0.92 1.35 0 . 8 9 1.42 0.92 1.42 0 . 8 9 1.35 1.23 1.65 0 . 8 9 1 .25 0 .89 1.42 1.00 1 .27 1.04 1 .39 1.00 1 .30 0.92 1.27 1.00 1.27 0.92 1.30 0 . 9 2 1.30 0.92 1.30 0,96 1 , 3 9 0.89 1.42 0.96 1.46 0.96 1.58 0 . 8 9 1.46 0 . 8 9 1.46 1.00 1.42 0.96 1 . 5 4 0 .92 1.42 1.04 1.23 0.85 1.42 0.85 1.46 0.85 1 .27 1.00 1.35 0.85 1.35 0.92 1 .39 1 .12 1 .27 53.48 79.86 University of Ghana http://ugspace.ug.edu.gh T a b l e 12. H e a d m e s u r e m e n t s o f w o r k e r s o f C. acvap i m e n s | s i n v a r i o u s c o l o n i e s N e s t 1 6 1 5 / 5 / 7 7 H W HL m H L 1 . 0 4 1.42 1.04 1 . 3 9 0.96 1 .54 0.98 1 . 5 4 0 . 8 9 1.35 0.98 1 . 5 4 0 . 8 9 1 .27 0 . 9 8 1 . 5 8 1.00 1.35 0.92 1 . 5 8 0.96 1 .62 0.85 1.35 0.96 1.42 0.85 1.27 1.12 1.39 1. 0 0 1 . 5 4 0.96 1.39 1 . 0 0 1 .27 1.12 1 . 3 9 0.85 1 . 5 4 0.96 1.46 0.85 1 .30 0.92 1.30 0 . 9 6 1 . 3 9 0 .89 1.42 0 . 9 6 1 . 4 6 0.96 1.46 1 .00 1 .46 1.00 1.46 O .85 1 . 2 3 0.85 1.42 0 . 9 6 1 .46 0.98 1.39 0 . 9 2 1 . 3 9 1.12 1.27 1 .00 1 . 4 2 0.81 1.27 0.92 1.27 0 .89 1.27 0 . 9 8 1.30 0.96 1.46 0.85 1 .30 0 .89 1.35 0.96 1 . 5 4 0 .96 1 . 5 4 0 . 9 2 1 . 3 9 0.92 1.46 1.12 1.46 0 .96 1.27 0. 9 2 1 . 5 4 0 . 8 9 1.27 0. 9 2 1 . 3 9 0. 8 9 1.46 0 .96 1.62 0 . 9 2 1 .39 0 . 9 2 1 , 3 9 0.96 1.65 0.92 1.42 53.84 81.34 University of Ghana http://ugspace.ug.edu.gh Table 13* Head mesurement of workers in various colonies of C. acvapimemsis Nest 17 30/5/77 H W H L m H L 0.96 1.42 0.96 1.46- 1 . 0 4 1.55 1.04 1 . 4 2 0 . 8 9 1.55 0.96 1 .62 0.96 1 .46 0 . 8 9 1.55 0 . 8 9 1 .25 CJ. 89 1 .42 1.00 1.42 1.00 1.35 0.92 1.42 0.92 1 . 5 4 0.96 1 . 5 4 1112 1 . 5 4 0 . 8 9 1. 4 2 0.96 1.42 1 .12 1.65 0.81 1 .27 1 . 0 4 1.46 1.04 1.46 1 .04 1.55 O .85 1 .27 0.81 1.27 0 .89 1.46 0.85 1 . 5 4 0 . 8 9 1 . 5 4 0.96 1.55 0 .96 1.62 1 .00 1.27 1.00 1. 3 0 1. 12 1.50 0 . 9 6 1. 5 9 1.25 1.46 0.85 1 . 4 2 0.96 1. 5 4 0 .96 1.55 0.92 1. 5 4 0.92 1,46 0.96 1.42 0.81 1.27 1.00 1.42 1.00 1 . 5 4 1 .04 1.55 1 . 0 4 1.59 0 .96 1.59 1 . 0 4 1.25 1.00 1.27 1 .04 1 . 5 4 1 . 0 4 1.27 1.00 1. 3 9 0.92 1.50 0.85 1.42 0.85 1 . 5 4 0 . 8 9 1 . 5 9 0 . 9 6 1.39 0 .92 1 .27 ' 55.77 82.38 University of Ghana http://ugspace.ug.edu.gh Table 14 Diurnal activity Faltem of C. r,cvnrdmoi:\s±3 on Z'h.hz Time Temp, Temp. Em. Leav. Entr, Bat: of day Air °C nest °c cif/ - nest nest Fooc 6 a.m. 23.0 21 .0 99 18 2 0 7 " 29.0 23.0 98 10 7 3 8 » 29.5 2C.0 90 7 15 6 9 " 27.5 30 .0 70 15 16 11 10 11 27.0 32.0 61 16 18 9 11 « 27.5 35.0 57 1 Q 0 12 Foon 54 • 5 !C.3 :o 18 V : 0 1 -%r . z\.o 41.0 ;o 16 6 4 2 ” 3U5 41.0 5f 6 4 2 3 » 33.4 3e.o 64 12 16 7 4 " 29.0 36.0 73 7 15 9 5 ,f 25.0 29.0 89 4 12 10 6 " 24.5 25.0 95 1 1t 4 Temr'erature recordin'; T*xas done by a thermometer. University of Ghana http://ugspace.ug.edu.gh Tablo 1 Diurnal activity Pattern of C. acYapimensls on 17/9/76 Time Temp. Temp. Hum. Leav. Eiitr. Efotr. Tjllh of M r nest £ nest nest food day °G °c 6 a .cu 22.5 21.5 98 12 3 0 7 ■ 26.0 24.5 94 6 4 1 8 " 28.0 28.0 b4 12 i . ’ -- 9 <« 2'. i.O 27,0 7'-> 17 14 11: 0 ooi- O a tfl 71 10 7 1 ■J !J 28o0 36.0 58 13 10 4 2 Noon 31.0 41.0 64 14 11 0 p.m. 31 o5 37.0 57 13 10 3 > ft 30.5 40.0 60 4 3 0 * " 30.0 35.0 70 9 13 6 u 28.0 32.0 82 8 17 8 ; » ■ 26.0 32.0 90 6 15 6 »» 23.0 24.0 95 13 14 2 Temperature recording w.qS done by a thermometer. University of Ghana http://ugspace.ug.edu.gh Table 16. Diurnal activity Toaltem of C . acvarlmongis on 1/10/7G Time of day Temp. Air °0 Temp. nest °C Hum. Leav . nost JSntr. 213 s t Entr.with food 6 a «m 0 25.0 24.0 99 3 3 0 7 " 25.0 24.0 97 0 1 1 8 " 25*0 25.0 87 2 2 1 9 " 27.0 25.0 31 0 4 2 10 " 30*0 30.0 69 1 3 1 11 « 32.0 29.0 62 2 1 1 12 I'Toon 32.0 29.5 54 0 0 0 1 p.m. 32.0 29.5 54 0 1 0 2 » 32.0 31.0 51 1 0 0 3 " 29.5 32.5 72 0 0e- 0 4 " 28.0 30.0 00 4 10 Ai 5 1 25.0 27.0 91 7 15 r7t 6 » 24.0 27.0 95 4 0 4 University of Ghana http://ugspace.ug.edu.gh Table 17. Divirn-l activity r»n>tem of C. CTa"imon3i^ on” I5/1 0 /7 : lime TQDlp, Ttimp* Evuaa j.G,?V. Entr. Eg tr* with of day Air °C nest °C $ n .1: t 113 st food 6 a.m. 22*0 25.0 1 3 0 7 " 24*0 27.0 95 2 4 1 8 « 30.0 31.5 08 0 1 1 9 « 32.0 33.0 75 1 0 0 10 « 31*0 34*0 70 t 2 t 11 « 31 *5 33.0 60 0 0 0 12 Foon 31*0 31.5 49 0 0 0 t p.m. 31 .0 32.0 70 2 1 1 2 » 30.0 31.0 70 5 AT 0 it 30*0 35.0 85 3 5 1 4 n 29.0 34.0 90 4 7 4 5 « 28*0 30 oO 94 4 6 3 6 « 25.5 28,0 95 1.9 10 f University of Ghana http://ugspace.ug.edu.gh Table 18* Diurnal activity/ paltern of G. -acvar>iinen3is on 29/10/7S Time Temp, Temp. Htmtm Leav. Sntr. J3ntr.tfd.th of day Air °C nest °c t/v ne3 t nest food 6 a.m. 23.0 25.5 98 1 4 0 7 " 28.0 29.5 94 2 4 2 8 » 28.5 30.5 ■34 3 2 1 9 " 29.5 32.0 70 0 1 0 10 ,! 30.5 34.0 71 5 2 1 11 " 30.0 34.0 58 4 0 0 12 Noon 31.5 35.0 64 2 0 0 1 p.m. 31.0 35.0 56 3‘ 1 0 p «» 31.0 34.0 54 3 0 0 5 ■ Vj ! V>J « O 33.0 64 4 6 T 4 " 30.0 33.0 81; 6 4 3 5 ,f 26.5 28.5 91 7 5 2 6 « 25.0 25.0 96 18 16 12 University of Ghana http://ugspace.ug.edu.gh Table 19. Diurnal activity on pattern 12/11/76 of C.acvaxiiaa-isx 3 Time Temp. 'I'emp. Hum. Leav. Entr. Sntr.iri ’;1i of d-ajr Air °C nest cc 7° nest nc\st food 6 a. 10.. 21.5 22.0 98 0 0 0 7 " 23*5 24.5 92 0 0 0 8 H 27.0 25.0 91 3 1 0 9 " 30 .0 26 *0 80 11 7 2 10 « 28,0 26.0 80 7 10 4 n » 28.0 26.5 72 14 6 1 12 Noon 30.5 27.0 69 18 18 4 1 31.5 27*0 70 13 15 3 2 » 33*0 28.0 61 12 14 12 3 " 34.0 28.0 61 8 *71 2 4 11 32«9 28.0 82 1 1 t 5 " 26.5 27.0 92 4 Oc. 1 C 11 24.5 26.0 98 W 21 11 University of Ghana http://ugspace.ug.edu.gh Diurnal activity partem of 0.acvat>lmensis on 26/11 /7o Table 20. Time Temp. Temp. Hum# Leav* Sntr, Sntr of Air nest V° nect nest foo day °0 °C 6 a.m, 26.7 26.7 97 2 1 0 7 " 26.7 26.7 94 0 0 0 8 " 29.2 23.3 91 10 9 2 9 ,f 29.2 29.4 83 52 50 24 10 " 31.7 30.8 84 81 00 30 11 « 32,2 31.4 71 35 55 16 12 H oon 31 o7 29.7 60 34 25 12 t Poltl* 31.4 30.3 61 21 22 6 2 ” 32.2 30.6 64 37 39 20 3 " 31.7 29.2 72 22 1b 5 4 " 30.0 27.8 81 23 1? 4 5 " 28.3 27.2 89 21 17 4 6 « 28.0 26.4 91 36 39 14 Temperature was recorded mtb a ©ecord: nri machine* ■r.th University of Ghana http://ugspace.ug.edu.gh Diurnal activity pattern of 0.acvaT>iiyy?nsjs on tO/l 2/76 Table 21. Time Tgmp. Temp. Hum* Leav. Entr. Entr .-with of day Xir °0 m st °c i nest n st food 6 a.m» 23.5 21.0 95 9 5 0 7 " 26.0 24.0 84 8 6 4 S 1 27.0 24.0 81 33 31 11 9 11 27.0 24.0 76 38 50 8 to " 29.5 25.0 73 31 34 13 11 n 31.5 26.5 63 7 7 3 1 2 Noon 31.5 26.5 55 29 '*'3 8 t pan. 30.5 26.5 50 12 14 10 2 " 30.0 27*0 53 5 6 0 3 ■ 29.0 27.0 62 4 14 1 4 M 29.0 27.0 71. 5 15 3 5 " 28.5 27.5 dO 10 20 12 6 » 27.0 27.0 90 15 18 0 111 temperature recoodin-;; dvr in;; activity paltern were recorded us in;;; the .eecordin0: machine. University of Ghana http://ugspace.ug.edu.gh Diurnal activity paltern, of G.acropiraenslo on 24/12/76 Table 22. Time of day Temp# Air °c Temp. nest °C Hum. J! Leav. nest Sn':r. nest Entr.with food 6 a.m. 26.0 24.0 91 15 7 4 7 » 26.5 25 #0 31 12 8 6 6 « 26.5 25*0 79 17 11 6 9 " 27.5 26.0 71 27 13 7 10 ,f 28.0 26.0 70 22 18 12 11 w 28.5 26.0 61 20 12 4 12 Noon 3®.0 27.5 52 18 24 18 1p*Kl© 31.0 27.0 47 27 12 6 2 w 31.5 27.0 52 16 22 16 3 « 31.5 27.5 53 5 7 6 4 11 30.5 27.5 73 6 12 S 5 1 29 .5 26.5 79 S 6 4 6 " 29.5 26.5 30 29 21. 16 University of Ghana http://ugspace.ug.edu.gh Table 23. D iu rn a l a c t i v i t y p a t te rn o f C.acvaulmensis on 14/1/77 Time of day Temp# Air °c Temp, nest °C Hum. % Leave nest Ehtr. nest Entr.with food & a#m0 24.0 21.5 97 7 9 5 7 11 25.0 23.5 91 15 8 4 8 » 28.0 25.5 87 24 7 6 9 M 29.0 27.5 80 12 15 11 10 " 31.0 30.0 74 19 23 12 11 « 31.0 30.0 73 36 14 12 12 Nooa 31.0 30.5 69 20 29 24 1 p.m. 32.4 30.5 65 30 22 18 2 " 32.0 31.0 59 12 17 11 3 tt 32.0 31.0 59 17 3 1 4 w 31.5 30 .0 62 12 3 2 5 " 31.5 30.0 62 12 3 Z 6 ” 28 .5 28.5 78 25 14 0 University of Ghana http://ugspace.ug.edu.gh on 28/1/77 Table 2 4o Diurnal activity pattern of G.acvapimensis Time of day Temp. Air °c Temp. nest °C Hum. % Leav. nest Entr. nest Entr,with food 6 a«m* 22.5 21.0 95 7 9 6 7 ti 24.5 21.0 91 11 12 8 8 ti 26.5 22.5 87 19 11 7 9 it 26.0 23.0 81 24 20 17 10 11 28 .0 23.0 76 29 18 14 11 11 29.5 25.0 71 12 5 3 12 Noon 29.5 26.0 65 17 11 9 1 p.m* 31.5 26.0 61 11 13 7 2 it 30.5 26.5 59 5 8 3 3 11 30.5 26.5 59 12 5 4 4 it 28.0 27.0 63 8 24 13 5 tt 28.0 27.0 65 9 15 12 6 it 27.5 28.0 69 41 37 14 University of Ghana http://ugspace.ug.edu.gh Table 25# Diurnal activity pattern of C.acvapimensis Time of day Temp. Air °c on Temp. nest °C 11/2/77 Hum. % Leav. nest Entr, nest Entr.with food 6 a»nu 21.5 22.5 97 12 10 2 7 ” 23.4 23.0 96 18 1 8 « 23.5 23.5 82 4 2 1 9 ,f 24.0 23.5 80 7 3 1 10 11 25.0 26.0 73 3 1 1 H 11 25.0 26.0 67 8 4 2 12 Noon 27.5 26.0 60 5 2 1 i p.m. 31.5 26.0 57 2 8 2 ” 32.5 25.0 53 2 7 5 3 " 33.5 25.0 54 1 5 3 4 11 33.0 25.0 61 17 4 5 11 30.1 24.5 72 8 7 3 6 « 29.5 24.5 84 9 1 1 Harmattan period. University of Ghana http://ugspace.ug.edu.gh Table 26 • Diurnal acrfcity pattern of Camp, acvapimensis on 25/ 2 /7 7 Time Temp. Temp. Him. Leav. Ehtr Entr of Air nest % nest nest ■with day °C °G food 6 a.m. 27.8 28.3 91 15 7 2 7 27.8 28.3 91 11 19 3 8 31.0 28.5 89 25 12 3 9 30.2 30.0 81 39 36 29 0 31.8 30.6 77 60 42 23 1 34.6 30.0 73 12 19 0 2 34.6 32.2 68 17 27 18 1 p.m. 33.9 33.4 65 29 21 18 2 30.0 32.2 72 88 35 14 3 28.9 28.5 85 12 15 4 4 27.8 28.9 87 15 7 3 5 27.8 28.5 91 12 5 ■12. 6 26.8 27.8 96 5 19 11 Increased activity due to a broken term tartan. University of Ghana http://ugspace.ug.edu.gh Table 2 7. Diurnal activity pattern of C»acvapimensis on 11/3/77 Time Temp® Temp. Hum* Leav. Sntr, 3ntr% of Air nest % nest nest wit?: day °c °c food 6 a0nio 30.6 29.5 97 5 7 2 7 ■ 30.6 30.0 87 12 11 9 8 " 33.0 33.0 78 10 18 12 9 " 35.5 31.6 77 15 23 15 10 " 32.2 33.4 68 6 5 2 11 u 34.6 32.2 65 2 1 0 12 Noon 32.2 33.9 63 0 0 0 1 p.9. 35.6 31.6 61 0 0 0 2 " 35.0 32.0 62 0 0 0 3 " 35‘.0 30.6 90 59 75 34 4 " 35.0 30 .0 88 40 25 21 5 " 35.6 30.6 88 40 16 8 6 " 31.1 28.0 89 10 21 19 The day was cloudy and there was a heavy downpour at 12 noon which stopped at half past University of Ghana http://ugspace.ug.edu.gh Table 2 8• Diurnal activity pattern of C«acvapimensis on 25/3/77 Time of day Temp0 Air °C Temp* nest 0 c Hum. 0/0 Leav. neat Entr* nest Entr*with food 6 a*m0 24.0 22.0 97 4 2 0 7 " 26,0 25.0 92 7 3 0 8 " 26.5 26.0 89 9 1 1 9 " 26.5 26.0 85 TO 5 0 10 " 28.0 26.0 81 3 1 1 Hi " 30.0 28.0 75 11 3 0 12 Noon 32.0 28.5 68 0 0 0 1 p.m. 33.0 31.0 71 2 4 1 2 " 33.9 31.5 73 2 7 1 3 " 32.6 31.5 79 2 12 3 4 " 31.6 29.5 82 4 8 2 5 " 30.6 29.0 87 3 7 1 6 " 30.6 29.0 91 9 1 0 University of Ghana http://ugspace.ug.edu.gh on 8/4/77 Table 29* Diurnal activity pattern of C«acvapimensis Time o f day Temp. Air °C Temp. nest °C Hum* % Leav. nest Entr. nest Ehtr.with food 6 ct.IHo 26.0 26.7 98 12 8 6 7 " 26.0 26.7 92 15 12 2 8 " 28.2 26.3 91 7 15 12 9 " 28.2 29.4 86 9 8 1 10 " 31.4 30.8 85 18 9 6 11: " 31.2 31.4 81 12 3 1 12 Noon 31.7 29.7 79 4 2 1 1 p.m. 31.8 30.3 79 7 7 3 2 • 32.2 31.6 83 3 5 4 3 " 31.7 29.0 87 2 3 2 4 " 30.1 26.8 92 1 9 0 5 " 28.3 26o8 95 5 4 1 6 " 28.0 26,3 97 7 5 2 University of Ghana http://ugspace.ug.edu.gh Table 30* D iu rn a l a c t iv it y p a tte rn o f C*acvapim ensis on 2 2 /4 /77 Time of day Temp* Air °0 Temp. nest °C Hum. % Leav. nest Entr. nest Eh tr. with food 6 a® m* 23.4 21.4 97 4 2 0 7 11 26 .4 23.5 95 9 6 0 8 " 28.0 23.5 91 1 4 2 9 11 28.0 23.5 86 18 9 4 10 " 30.6 25.0 82 34 14 9 11 11 31.6 29.0 71 1 0 0 12 Noon 31.6 29.0 71 1 © 0 1 p.m* 32.4 29.0 75 4 3 0 2 ” 33.0 29.0 82 7 12 6 3 " 33.0 29.5 86 3 15 9 4 " 31.6 29.0 91 1 8 3 5 11 31.6 28.5 92 5 4 0 6 " 31.0 2B.5 95 1 2 1 Light intensity was high but the larvae and pupae were many yet activity izpas low© University of Ghana http://ugspace.ug.edu.gh Table 31« D iu rn a l a c t iv it y p a tte rn o f C>acvapim ensis on 6/5/77 Time of day Temp. Air °G Temp. nest °C Hum. % Leav. nest Ebtr. nest Eh t r . with food 6 a»m0 27.4 24.0 98 12 15 3 7 tt 27.4 24.0 95 34 21 11 8 tt 30 .0 26.4 93 11 22 17 9 it 31.0 26.4 91 32 15 7 10 fi 31.8 28.0 87 12 19 9 11 it 33.6 28.0 86 11 12 T1 12 Noon 33.6 29.5 83 6 9 7 1 p .m . 33.6 29.5 89 2 12 6 2 tt 31.6 30.0 81 14 24 7 3 tt 28.6 30.0 87 26 5 0 4 tt 28.6 28.0 90 3 2 1 5 tt 27.8 28.0 90 9 3 2 6 tt 26.0 28.0 93 12 4 1 Jl very sunny day bui ants were very active due to a, food source 0 University of Ghana http://ugspace.ug.edu.gh on 20/5/77 Table 32. D iu rn a l a c t iv it y p a tte rn o f C .aavapim ensis Time of day Temp# Air °C Temp. neat °C Hum. % Leav. nest Entr. nest Eh tr. with food 6 a«m« 28.3 26.7 97 12 16 3 7 ti 29.2 28.0 95 19 12 5 8 u 27.2 27.2 91 13 18 7 9 n 26.7 27.2 87 29 19 9 10 n 28.0 29.2 83 16 13 2 11 it 28.3 29.2 76 23 19 0 12 Noon 29.2 29.4 78 36 28 3 p.m. 28.0 29.5 79 15 34 2 2 ti 30.6 27.2 83 22 38 15 3 it 29.2 27.2 89 8 17 7 4 it 31.4 28.3 91 23 12 9 5 ti 33.3 30.0 94 12 18 8 6 ti 33.9 31.7 94 34 22 7 University of Ghana http://ugspace.ug.edu.gh on 3/6/77 Table 33o D iu rn a l a c t iv it y p a tte rn o f 0«acvapimens i s Time of day Temp. Air °G Temp. nest °c Hum. % leav. nest Entr. neat Entr.with food 6 aom# 26.1 26.7 98 9 7 3 7 26.7 28.0 95 13 14 7 8 If 28.3 28.3 92 19 12 9 9 It 27.2 28.3 89 7 7 2 10 11 29.4 29.2 81 8 12 0 11 II 30.0 29.4 90 14 16 2 12 Noon 27.0 28.0 96 26 36 7 1 p.m. 26.7 28.0 95 36 45 11 2 it 27.2 28.0 95 40 45 19 3 ii 29.2 30.0 86 15 15 5 4 it 30.6 32.8 85 5 11 Aft 5 ?t 32.2 31.4 92 6 12 6 6 i i 21.2 28.3 97 23 29 8 University of Ghana http://ugspace.ug.edu.gh Table 34- Diurnal activity pattern of C«acvapimensis Time of day Temp# Air °c on Temp. nest °C 17/6/77 Hum. % Leav. nest Sntr. nest Eh tr. with food 6 a.m. 24.0 23.9 99 11 12 8 7 it 24.0 23.4 98 15 11 0 8 tt 23.8 23.4 98 24 25 12 9 ti 23.9 23.3 98 17 12 9 10 ft 23.9 23.4 99 19 11 3 11 tt 24.0 23.5 98 23 8 7 12 Noon 24.0 23.5 99 23 7 4 1 p .m . 24.0 23.0 99 15 19 16 2 tt 24.0 23.0 99 23 13 12 3 tt 23.9 23.0 99 32 17 11 4 it 22.9 22.0 100 17 24 1.1 5 it 23.0 22.5 100 18 11 7 6 23.1 22.5 100 15 12 2. Leavae were many, the day was cloudy and misty with drizziling* University of Ghana http://ugspace.ug.edu.gh Table 35* Diurnal activity pattern of C«acvapimensis on 1/7/77 Time of day Temp. Air °C Temp. nest °C Htmu % Leav. nest Entx. nest 2ntr.with food 6 a.m. 25.4 22.3 100 17 13 11 7 " 25.3 23.4 96 32 21 8 8 " 25.4 23.5 92 12 15 12 9 " 27.3 23.5 89 19 32 21 10 " 28.0 24.0 78 25 17 11 11 tt 28.9 26.1 72 13 12 5 12 " 28.9 26.1 74 17 29 17 1 p.m. 29.5 26.5 73 29 18 21 2 " 29.8 26.9 78 43 17 12 3 " 30.1 26.9 83 16 23 11 4 " 29.1 27.0 82 32 39 19 5 rt 28.4 27.0 89 17 28 20 6 " 28.0 27.1 90 19 12 1 University of Ghana http://ugspace.ug.edu.gh Table 3 6. Diurnal activity pattern of £.aovapimensis on 15/7/77 Time of day Temp 0 Air °c Temp. nest °C Hum, % LeaVo nest Entr, nest 2a tr,with food 6 aoDle 27.4 26.3 99 15 30 17 7 1 27.9 27.1 97 21 12 4 8 « 27.9 27.2 93 32 16 12 9 " 30.2 28.4 87 17 39 13 10 11 31.4 28.9 83 23 12 1 11 " 31.4 29.0 87 19 11 1 12. 11 32.6 29.1 81 45 14 5 1 p.m. 32.4 29.0 78 28 33 12 2 " 31.0 29.3 75 32 17 13 3 " 30.4 28.7 77 12 29 15 4 11 30.0 28.5 82 19 11 1 5 " 29.5 28.8 89 23 12 7 6 « 29.1 28.8 91 32 17 3 University of Ghana http://ugspace.ug.edu.gh ^iurnal activity pattern of G.acvapiraensis on 29/7/77 Time Temp. Temp. Hum. Leav. Entr. Entr.with of Air nest % nest nest food day °C °C Table 37. 6 a.m. 27.4 21.4 99 26 11 7 7 " 27.0 23.5 97 12 29 18 8 " 30.3 25.0 93 17 16 12 9 " 30.5 25.0 89 31 42 23 10 11 30.5 29.5 86 29 11 4 11 " 31.5 29.4 87 16 29 6 12 .. 32.0 29.0 83 34 17 8 1 p.m. 32.0 29.0 81 12 28 13 2 " 31.0 28.5 87 18 32 15 3 " 31.0 29.0 89 25 17 11 4 " 29.0 29.5 91 12 11 10 5 " 29.0 29.5 93 17 23 12 6 " 29.0 29.5 95 19 24 14 University of Ghana http://ugspace.ug.edu.gh Table 380 Diurnal activity pattern of C.acvapimensis on 1.2/8/77 Time of day Temp* Air °c Temp. nest °c Hum* % Leav. nest Entr. nest JShtr.vrith food 6 a*m* 22.8 22.8 98 12 17 4 7 ti 22*9 23.0 96 19 11 7 8 ti 23.0 23.3 92 29 12 8 9 i i 23.5 23.9 91 39 26 12 10 tt 23.6 23.9 87 54 33 26 11 w 24.0 24.6 86 41 51 39 12 Noon 24.3 24.8 81 44 53 34 1 p.m. 24.9 24.8 79 37 15 11 2 tt 25.0 26.5 78 39 46 29 3 tt 25.0 26.5 83 60 51 41 4 ti 24.9 26.0 86 32 43 32 5 it 24.6 26.0 91 12 17 7 6 tt 24.0 26.1 94 19 25 21 University of Ghana http://ugspace.ug.edu.gh Table 39. Diurnal activity pattern of C.acvapimensis on 26/8/77 Time of day Temp. Air'; °C Temp. nest °C Hum. % Leav. nest Entr. nest Entr.with food 6 a.m. 23.0 22.1 91 29 17 7 7 " 23.1 22.0 96 17 21 9 8 '• 23.1 22.0 96 34 18 15 9 " 23.4 22.0 96 43 23 12 10 " 23.3 22.3 97 27 12 1 11 " 23.4 22.3 97 28 11 0 12 " 23.5 22.4 98 12 17 15 1 p.m. 23.1 22.3 98 18 13 1©- 2 " 23.1 22.1 99 22 33 8 3 " 23.6 22.3 98 34 12 9 4 " 23.0 22.3 99 16 15 7 5 " 23.0 22.4 99 11 21 14 6 " 22.8 22.4 100 30 38 12 University of Ghana http://ugspace.ug.edu.gh Time Temp* ' Temp* Huni. Leav* Sntr* 3ntr*^ith of Air nest % nest nest food Day °C °c Table 40: Noctural activity pattern of C*acvapimensis on 14/10/7o 6 a.m 2(5.4 25.5 96 35 45 20 7 " 27.8 26.1 94 25 32 10 8 " 27.5 26.1 93 3 13 4 9 " 27.8 26.1 95 £5 27 8 10 " 27.5 24.7 95 39 43 6 11 " 27.2 23.9 96 65 59 2 12 noon 2S.7 23.9 98 18 10 8 1 P.m 26.4 23.6 92 17 13 6 2 " 26.1 23.3 93 30 39 7 3 " 25.8 23.3 94 27 36 1 4 •• 25.5 23.1 97 13 33 17 5 " 25.5 22.8 98 16 24 12 6 " 25.3 23.1 97 2 1 0 University of Ghana http://ugspace.ug.edu.gh Table 41 i No choral activity pattern of 0#.aovap.ij-\ensi3 on 28/10/7 6 Tine of day Temp* Air °G Tenrp. nest °C Humi. cr/° Leav. nest Ent. nest Ent.with food 6 a*m 25.0 24.5 91 36 39 14 7 " 26.0 23.6 96 19 20 6 8 " 26.5 23.6 95 5 6 0 C) n 26.5 23.6 96 26 3 0 10 » 27.0 23.6 96 26 31 5 11 " 27.4 23.0 97 57 53 6 12 noon 27.4 24.0 96 10 12 *O 1 p.m 26.5 24.0 94 11 15 11 2 " 26.3 23.0 94 42 36 3 3 " 25.4 23.0 95 29 32 3 4 " 25.3 23.5 96 25 36 18 5 11 25.3 23.6 98 21 25 15 6 * 25.3 23.6 97 7 6 0 University of Ghana http://ugspace.ug.edu.gh Table 42: Noctural activity pattern of 0 »acvajoiivierisis on 11/11/76 Time of Day 'Temp. Air °C Temp. nest °C Humi­ dity So leav nest 3nt. nest Iht.w: food 6 a*m 26.0 26.0 94 28 33 15 7 H 27.1 25.5 94 27 30 11 8 " 27.5 25.5 96 4 8 3 9 " 27.5 25.5 94 64 61 15 10 n 27.8 25.0 95 43 42 •7t 11 " 26.7 24.5 96 7 11 4 12 noon 26.0 24.5 96 72 67 10 1 p*m 26.1 24.5 92 51 50 11 2 » 25.3 23.1 92 75 67 4 3 • 25.4 23.0 93 68 62- 3 4 » 25.5 23.0 97 49 4 6 15 5 " 25.0 23,0 98 32 15 11 6 " 25.0 23.0 98 10 8 1 University of Ghana http://ugspace.ug.edu.gh Table 43 : Nootural activity pattern of C.acvaDimensis on 25/11/76 Time Temp. Temp. Humi­ Le^v. 3nt. Sit. of Air nest dity nest nest Tilth Day °c °C o 'S'* food 6 a.m 27.0 25.0 97 20 29 12 7 " 28.0 24.5 96 21 27 OO 8 « 27.0 25.0 97 12 15 1 9 11 26,5 25.0 96 60 56 0 10 " 26.5 24.5 98 41 50 2 11 " 26.0 23.1 97 5 0> 1 12 Noon 26.3 23.5 97 61 O 'J 2 1 p*m 26.0 23.5 95 47 52 8 2 ?? 25.4 23.1 95 70 63 71 3 w 25.8 23.1 96 40 25 1 4 » 25.5 23.0 97 49 40 11 5 ” 25.0 23.0 93 32 26 12 6 * 25.0 23-0 97 11 4 0 University of Ghana http://ugspace.ug.edu.gh Table 44: Noctural activity pattern of C,aov^.i^en-ls on 9/12/7 5 Time Temp* Temp. Humi­ Leav. Sntr. a of Air nest dity nest nest Day °c °C c' f c 6 a#m 26.4 24.5 100 29 32 24 7 * 25.4 24.0 98 20 13 14 8 " 25.0 23.0 97 17 24 21 0 .» 25.0 23.0 98 48 52 41 10 1 24.5 22.5 96 46 41 1? 11 " 24.5 22.5 96 4 6 4 12 Noon 24.5 21.8 98 54 21 12 1 p.m 24.0 21.5 97 40 49 33 2 « 23.5 21.5 95 62 67 41 3 " 23.5 21.5 97 47 41 32 4 ,f 24.5 22.0 96 45 12 7 5 " 24.0 22.0 98 33 48 42 6 ” 24.0 22.0 98 19 22 12 University of Ghana http://ugspace.ug.edu.gh Table 4 5: Nootural activity pattern of HLaovapimensls on 23/1 2 /7 6 Time of Day Temp. Air °c Temp. nest °C Humi­ dity % Leav. nest Bntr. nest Sntr. with. food 6 a*m 26.0 24.0 98 36 38 32 7 11 26.0 24.0 97 24 27 21 8 " 25.5 23.0 97 22 18 16 25.5 23.0 96 31 17 8 10 " 23.0 22.0 97 42 54 50 11 * 22.5 21.0 98 12 17 12 12 Noon 22.0 21.5 98 59 50 47 1 p.m 22.0 21.5 98 31 37 12 2 " 22.0 21.5 97 52 49 37 3 tt 22.5 21.5 97 44 51 42 4 rt 22.5 81.5 97 22 17 4 5 " 23.0 21.0 98 30 28 21 6 « 23.0 21.0 99 12 17 8 University of Ghana http://ugspace.ug.edu.gh Time Temp# Temp# Kumi— Leav* 3ntr* X2n.tr* of air nest dity nest nest v/ith Day °C ° C % food Table 46s Nootural avtivity pattern of O.aovapimeasis on 6/1/77 6 a*m 0• I"-CM 27.8 99 42 27 21 7 « 26.2 27.8 98 30 31 25 8 t! 25.0 27.8 98 29 22 18 9 »» 25.0 28.5 97 17 29 27 10 H 25.0 26,8 98 57 49 41 11 n 24.4 27.3 98 18 7 3 12 Noon 23.9 26.8 98 51 53 48 1 p*m 23.9 26.8 98 39 41 30 2 ti 23.4 26.8 97 48 40 3'6 3 « 23.6 26.6 97 36 21 8 4 tt 23.4 26.2 97 18 14 12 5 11 23.4 25.6 98 42 26 17 6 w 25.0 26.2 98 4 18 12 University of Ghana http://ugspace.ug.edu.gh Table 47 s Hoctural activity pattern of Oaacva~oimensis on 20/1/77 Time of Day Temp. Air °c Temp, nest °c Humi­ dity % Leav. nest 2rj.tr. neat Etr';r. ■with food 6 a*m 26.4 25.5 95 18 4 2 7 " 27.8 26.1 97 12 7 5 8 ff 27.5 26.1 97 1 2 1 9 " 27.8 24.7 98 5 3 3 10 " 27.5 23.9 98 1 7 0 11 ” 27.2 23.6 96 2 7 4 12 Noon 26.7 23.3 97 2 4 3 1 p*m 26. 4, 23.3 96 0 1 0 2 » 26.1 23.1 96 8 5 ~yJ 3 " 25.8 23.1 9 6 16 14 nI 4 " 25.3 23.1 98 19 16 9 5 " 25.3 23.1 98 15 8 6 6 * 25.8 24.4 100 7 11 4 University of Ghana http://ugspace.ug.edu.gh Table 48: Noctural activity pattern of C.acvapinensis on 4/2/77 Time of Bay Temp. Air 0 c Temp. nest °C Htuni- dity 0/ Leav. hast Zfatr. nest Bn ;r. rrith food 6 a<,m 27.4 24-3 99 15 17 4 7 ,f 27.4 24.7 97 29 13 9 8 " 21.2 24.6 97 19 52 12 9 / 26.5 24.3. 97 12 34 13 10 " 26.4 24.2 95 17 29 17 11 " 26.8 24.0 96 22 31 15 12 Ho on 25.4 24.2 96 73 59 41 1 p#m 26.2 24.1 97 16 19 12 2 11 25.7 24.0 98 51 29 7 3 " 25.4 24.0 98 48 21 3 4 ” 25.5 24.0 98 39 17 9 5 ” 26.0 23.6 97 17 12 4 6 w 25.8 23.3 96 15 8 1 University of Ghana http://ugspace.ug.edu.gh Table 49s Koctural activity pattern of C.aovapim&nais on 18/2/77 Time of Day Temp. Air °c Temp. nest °C Humi­ dity Leav. nest Sntr. nest Ihtr. vdtjr food 6 a#m 28.3 24.3 95 35 24 17 7 11 27.4 24.1 95 39 29 12 8 " 27.3 23.2 97 42 S3 43 9 ” 27.0 23.3 98 12 10 7 10 11 27.0 23.1 97 72 19 12 11 11 25.3 23.1 98 49 35 28 12 noon 25.0 23.4 97 31 63 19 1 p.m 25.0 23.4 93 27 18 4 2 11 21.3 22.8 97 9 19 11 3 " 24.7 22.9 98 18 34 12 4 ” 25.0 23.0 97 23 C>0 16 5 11 23.7 23.7 95 29 12 sO 6 w 25.7 23.7 98 12 18 S University of Ghana http://ugspace.ug.edu.gh Table 50: Nootural activity pattern of C.aovapijaensis on 4/3/77 Time Temp# llEtnp. Humi­ Leav. Sntr. Srxtr. of Air ne3t dity nest nest TTith Day °c °c 5° food. 6 a.m 27.8 26.8 98 43 22 12 7 " 27.8. 27.0 97 19 34 17 8 " 28.5 26,2 97 35 12 4 9 " 26.8 25.0 96 39 19 O 10 » 27.3 25.0 94 70 12 5 11 “ 26.8 24.4 97 19 39 11 12 Noon 26.8 23.4 97 72 12 3 1 p.m 26.8 23.9 98 21 29 16 2 " 26.6 23.9 98 28 04 32 3 " 26.2. 23.5 97 13 12 10 4 " 25.6 23.4 97 33 19 11 5 " 26.2 23.4 97 11 29 4 6 " 26.4 25.5 98 9 1.8 7 University of Ghana http://ugspace.ug.edu.gh Table 51: Noctural activity pattern of C«acvap 1mensis on 18/3/77 Time o f Day Temp * Air °c Temp. nest °C Humi­ dity o'/° Leav. nest 3n ;r. nest Sntr. Tith food 6 a*m 28.J 27.8 99 73 19 12 7 51 24.4 25.4 98 89 12 10 8 « 26.2 25.6 97 70 21 4 9 11 25.0 25.0 97 17 19 qy 10 26.2 25.0 97 22 34 18 11 !t 25.6 25I3 96 40 62 33 12 Noon 26.2 25.2 96 17 31 12 1 p.m 26.8 25.4 97 19 39 17 2 " 27.3 25.5 96 33 28 13 3 " 26.2 25.3 9S 39 17 Cl* 4 tf 26.2 25.6 98 12 19 3 5 * 26.2 25.6 93 17 15 9 6 » 25.6 25.4 99 33 20 5 University of Ghana http://ugspace.ug.edu.gh Table 52: Noctural activity pattern of C♦ acva£faensis on 2/4/77 Time of Day Temp# Air °c Temp. nest °G Humi- dity 55 Leav. nest Un '.r. nest Entr. with food 6 a«>m 29.4 27.8 99 17 18 9 7 " 25.3 25.2 99 34 21 12 8 " 25 .0 24.3 100 19 16 3 9 f 25.0 24.8 100 38 12 5 10 ” 2S.3 f \3 • CO 100 37 14 6 11 « 26.2 2£#8 99 45 52 18 12 Ho on 25.6 23.8 97 73 59 42 1 p.-m 25.6 24.0 98 12 17 3 2 » 25.4 24.0 98 18 39 12 3 " 25.4 25.0 98 27 25 17 4 " 25.4 25*0 98 5 3 1 5 11 26.2 25.0 98 19 22 13 6 " 25.6 25.0 97 8 4 1 University of Ghana http://ugspace.ug.edu.gh Table 53* Nootural activity pattern of C.acvap im ens is on 16/4/77 Time Temp. Temp. Humi­ Leav. Ifotr* Ihtr. of Air nest dity nest nest V/l b_n Day °c °C y~ 6 a*rn 28.7 26.2 98 6 12 2 7 ii 28.0 26.0 97 19 12 8 8 i? 26.2 25.0 95 12 16 12 9 it 25.6 25.0 93 34 23 11 10 tt 25.6 23.6 96 16 19 12 11 II 25.6 23.6 96 39 15 9 12 Noon 26.8 23.6 97 43 13 G 1 p.m 26.8 23.4 95 71 46 23 2 51 27.3 23.4 98 12 Jo ig 3 t t 27.3 25.0 qo ✓ ✓ 11 29 11 4 / 26.0 25.0 98 14 23 13 5 II 26.0 25.6 97 19 12 4 6 I t 25.6 25.0 00 23 17 /•0 University of Ghana http://ugspace.ug.edu.gh Table 54: Nootural activity pattern of C^ aovspliflaasl.s on 30/4/77 Time of Day Temp. Air °c Temp. nest °c Humi­ dity % Leav. nest Sntr. nest Sn ;r. with food 6 a«m 26.0 25.0 99 7 6 3 7 ft 26.4 25.0 98 9 12 4 8 n 26.4 26.0 98 15 5 2 9 t 27.0 26.0 97 32 13 5 10 n 27.0 26.0 97 32 12 5 11 tt 27.0 24.7 96 24 21 8 12 Noon 27.2 24.7 96 16 13 2 1 p*m 26 .4 23.3 97 18 12 5 2 tt 26.4 23.3 98 23 15 6 3 tt 25.5 23.6 98 29 7 2 4 tt 25.5 23.6 97 24 17 8 § tt 25.5 23.1 93 26 13 9 6 tt 26.4 23.1 99 27 13 6 University of Ghana http://ugspace.ug.edu.gh Table 55 Time of Day : Nocttiral activity pattern of on 14/5/77 Temp. Temp. Humi- Leav Air nest dity nest ®c °c % C&acvap imensis Sat. Sntr. nest with fool 6 a.m. 26.0 26.0 98 3 9 1 7 " 26.0 26.1 98 11 7 z 8 " 27.8 26.1 97 29 12 3 9 11 28.3 24.0 97 6 4 0 10 11 28.0 23.3 96 14 32 6 11 27.2 23.6 97 23 19 7 12 Noon 27.8 23.6 96 17 11 5 1 p*m 27.5 23.3 95 4 9 2 2 ” 27.2 23.6 95 15 9 2 3 " 27.3 23.1 96 19 14 5 4 " 26.4 23.1 97 17 7 2 5 " 26.4 24.4 97 29 16 3 6 " 26.4 24.9 97 13 11 4 University of Ghana http://ugspace.ug.edu.gh Table S£>: Hontiiral aotivitv nattem of C.acvapiraensis on 28/5/77 Time of o&y Temp. Air °c Temp. nest °C Humi­ dity o/ Leav. nest Sncr. nest Etrfcr. with food 6 a.m 21,8 26.1 99 16 24 12 7 " 28.0 26.1 97 28 36 11 8 " 28.0 26.1 93 33 29 13 9 " 27.8 26.4 94 29 27 17 10 " 27.8 26.1 93 51 63 34 11 " 27.5 26.1 97 27 15 4 12 noon 27.3 25.8 96 29 11 2 1 p.m 27.2 25.5 96 33 15 7 2 " 27.2 25.3 98 29 38 16 3 n 26.4 25.0 99 14 9 7 4 " 26.7 25.0 97 12 6 1 5 " 26.7 25.0 97 16 9 5 6 " 26.0 25*0 99 12 4 0 University of Ghana http://ugspace.ug.edu.gh Table 57* Nootural activity pattern of C.aoya'0iraensis on 11/6/77 Time of Day Temp. Air °C Temp. nest °C: Humi­ dity % Leav. nest Bnfcr. nest Entr. with food 6 a.m 22.4 21.9 98 12 4 0 7 " 22.6 29.7 98 7 12 0 8 " 22.5 21.5 96 3 5 1 9 " 22.3 20.9 99 2 14 1 10 ” 22.1 20.9 98 15 7 2 11 " 22.0 21.0 97 25 21 4 12 " 21.0 21 *0 97 17 12 2 1 p.m 21 .7 21.1 98 11 19 4 2 » 21.9 21.0 97 4 7 0 3 " 21.7 20.9 97 9 16 5 4 " 21.5 20 08 97 11 4 1 5 " 21.9 20.9 97 13 9 0 6 " 22.0 21.2 98 5 1 0 University of Ghana http://ugspace.ug.edu.gh Table 58: Nootural activity pattern of C♦ acvai-Anensis. on 25/5/77 Time of Day Temp * Air ° c Temp. nest I* Humi­ dity 0//« Leav. nest Sfotr. nest 6 a.m 21.9 22.5 99 5 2 7 u 22.9 23.9 99 7 1 8 11 22.5 24.2 99 12 3 9 n 22,1 23.8 98 11 0y 10 11 23.4 23.2 100 9 5 11 it 23.5 23.2 100 18 12 12 Noon 24.4 22.5 99 3 n1 1 p.m 24.1 22.5 99 2 r—2 2 it 24.5 22.0 100 5 15 3 it 23.9 22.0 99 17 12 4 tt 23.9 21.9 99 4 n t 5 it 23.9 21.9 99 O 9 6 « 23.9 21.9 100 1 3 Entr, v/itli food 0 0 0 4! 4 1 2 5 4 2 1 0 University of Ghana http://ugspace.ug.edu.gh Table 59s Noctural activity pattern of C*acvapimensis on 9/7/77 Time of DaJ Temp. Air °c Temp. nest °C Humi­ dity Leav, nest En.tr, nest 33h ;r. with food 6 a.m 22.4 22.5 99 7 5 1 7 " 22.9 23.9 98 9 19 5 8 '» 22.5 24.3 97 12 11 3 9 " 22.3 24.2 96 15 12 2 10 " 22.0 24.2 96 3 14 4 11 " 22.0 23.9 97 12 3 1 12 " 22.4 23.9 97 17 2 0 1 p*m 22.5 23.9 9S 3 5 o(— 2 » 22.5 22.6 97 9 16 3 3 " 22.6 22.9 97 8 9 2 4 w 22.4 23.4 97 7 12 4 5 " 22.4 23.4 98 16 4 1 6 w 22.9 23.6 99 2 3 0 University of Ghana http://ugspace.ug.edu.gh Table 6p: Noctural activity pattern of C.acvapisiensis. on 23/7/77 Time of Day Temp, Air °C Temp# nest °G Humi­ dity e' leav. nest 3ntr. nest Hhtr. with food 6 a*m 24.4 25.0 99 7 4 1 7 " 26,4 25.0 99 3 12 2 8 « 27.0 26.0 99 13 9 1 9 0 28.0 26.0 97 15 3 0 10 » 27.2 24.3 96 6 2 0 11 11 27.8 24.7 96 12 9 0 12 Noon 27.5 24.7 96 17 11 3 1 p.m 27.2 24.0 97 25 4 2 2 tf 27.3 23.6 98 19 12 3 3 " 26.4 23.4 99 4 15 4 4 w 26.4 23.4 100 17 12 3 5 11 26.4 23.2 99 5 9 1 6 w 26.4 23.1 99 7 3 0 University of Ghana http://ugspace.ug.edu.gh