Butterflies and moths (Lepidopterans) occur across continents and are a well-known and popular group of insects, enriching the lives of many people. They are a source of food and house hold income in many African countries, possess aesthetical beauty and therefore, an attraction for local and foreign tourists, some are important crop pests, at least in one or two of their life cycles,  and also play a critical role in the ecosystem.

 In Zambia, several hundred species of tropical and subtropical butterflies occur across the entire country. This great variety of butterflies is largely attributed to the country’s abundant and wide spread water resources, varying habitats which support a wide range of vegetation communities and diversity of plant species. This makes Zambia a suitable location or destination for the study, collection (permitted by DNPW) or simply outdoor pursuit of butterflies and moths alike. 

The delightful colour patterns of butterflies confer an inherent aesthetic appeal to both amateur and professional collector and visitor. This is obviously an undeveloped niche in nature based tourism which has the potential to contribute to the increased visitor retention period and satisfaction levels and hence to the growth of the nature based tourism subsector.

 [1] however, observed that butterflies among other members of the insect group despite their socio-economic and ecological importance are vulnerable to natural and anthropogenic factors. For instance, in their early stages of life, butterflies have to contend with weather changes, as well as natural enemies, wild fire, habitat loss and in some cases over collection by amateurs and professional collectors, agriculture, expanding industries, use of pesticides and pollution on one hand while the clearing of woody vegetation can reduce food plants (particularly nectar yielding plants) and hence butterfly populations. Without care, quite a number of non-harmful insects can be knowingly or unknowingly be exterminated. 

 Of critical concern now in the medium to long-term time scales is the survival of butterflies and other insects is the subsisting environmental impacts arising from climate change. Because of their sensitivity to changes in the environment, butterflies have been recognized as a useful biodiversity indicator group of tropical land-use systems because as stated above,  they are sensitive and react quickly to subtle changes in environmental and habitat conditions [2-7] Due to their short life cycle, narrow niches and relatively low mobility, they are more sensitive to land-cover and land-use changes than long-lived animals.

It is a known fact that butterfly activities are closely controlled by weather and many species are constrained by climate, usually occupying a small part of the range where host plants as food and cover subsist as acknowledged by [8] Butterflies also play a significant ecological role in agricultural landscapes,  are sensitive to changes in the environment, easy to see and study, and for this reason, we considered them to be suitable candidates for monitoring changes in habitat, biodiversity and environmental conditions as earlier reported by [2,9,10] including the impact of landscape and habitat management practices and disturbance regimes in terrestrial ecosystems [11,12]

Despite their smaller size, [13] Schmidt and[14], succinctly reported on their role in the recycling of nutrients (N, P, K) highly needed by crops thereby supporting agriculture (crops). In their larval stage, butterflies feed on leaves of several wild plants found in the agricultural systems and therefore release faeces that contain large amounts of nutrients [15] In addition, butterflies are food to birds and other predators and are hosts to several parasitoids that suppress crop pests [16,17] This important ecological role underscores the need to conserve butterflies in order to sustain the productivity of natural and agricultural landscapes. 

 In Zambia as is the case with several sub-Saharan African countries, butterflies are not the people’s choice, and most studies as earlier mentioned are skewed towards large and more conspicuous species such as mammals, reptiles and birds. It is indeed common knowledge that monitoring butterflies is not a priority as attention and policy direction of many nature based public institutions as well as NGOs is overwhelmingly tilted towards mega fauna such as elephant, rhino, lion and others. Because of this bias, butterflies remain relatively obscured and little studied, particularly with regard to their ecology, behaviour and functional role in farmland habitats both in natural areas and agricultural landscapes. [18,19] made similar observations that although butterflies made enormous contribution to pollination of plants including crops, there was no marching attention in terms of research in Africa  to have a full understanding of their ecology. To the contrary, knowledge of butterflies inhabiting farmland habitats is good in Mediterranean regions compared with the sub-Saharan Africa. The relative scarcity of data on tropical butterfly populations hampers the ability to effectively conserve them, particularly as pollinating agents [20] in agricultural systems.

1.1 Economic Importance of Butterflies and Moths 

Although this study was carried out specifically on butterflies in a National Park, its findings have a crosscutting and scientific application for the order Lepidoptera and socio – economic application in Zambia and many other countries in the SADC region where many species of caterpillars form an important source of protein. Of particular importance is the speckled yellow and white larva of the moth (Lepidoptera: Saturniidae which is distributed throughout the southern regions of Africa) and popularly called Mopane caterpillar. It is called mopane worm essentially because in Zimbabwe, Botswana and Namibia it feeds on the leaves of the mopane (Colophospermum mopane) tree, but a variant of it in northern Zambia depends on miombo trees (Brachystegia, Jubernardia, Isoberlinia and their associates) and it is called by a different name in local dialects (Mpika, Kasama, Mungwi, Senga Hill, Nsama and Mporokoso districts). The caterpillar is known to be a source of Iron and Zinc and is also high in protein (48.3%) similar to soybean [21] and is considered an important source of food for many people throughout this region of Africa. The adults, known as the emperor moth, are quite large with conspicuous eyespots on the hind wings (Figure 1a, b).

[21,22] for instance, showed the life cycle of Gonimbrasia belina (Lepidoptera: Saturniidae) and stressed that it was a nutritional food source rich in protein, fatty acids, and minerals and its popularity as food not only for human direct consumption but as a component of livestock feed in the southern region of Africa including Zambia (Figure 1a, b). 

Figure 1a: Life cycle of Imbrasia belina a) eggs, b larva –edible stage, c) pupa and d) adult female [22]

Figure 1b: Larva of the popular Imbrasia belina (Source: Favourite snack mopane worm, nydailynews.com)

 On the aspect of tourism, butterflies carry an undisputed aesthetic beauty different from large animals particularly in open landscapes such as the Kafue flats where watching butterflies is a priceless adventure. 

In conserving butterflies and moths therefore, we need to underscore the concern that climate change poses a huge threat to our butterflies, but that there are also ways to mitigate negative impacts and save them in our gardens, back yards, protected areas  and countryside in general [23] Butterflies have short life cycles and thus react quickly to environmental changes. Their limited dispersal ability, larval food plant specialization and close-reliance on the weather and climate make many butterfly species sensitive to fine-scale changes and therefore suitable indicators of habitat condition. Because, butterflies are good biological indicators of environmental change, holistic measures to help them will assist in conserving Zambia’s rich biodiversity.  Preserving the natural habitats in areas such as the Lochinvar National Park, which offers an array of large habitats with a mosaic of microclimatic conditions, can undoubtedly help sustain larger and more stable butterfly populations and communities for a much longer time than small areas particularly in isolated urban habitats. 

1.2 Impact of climate change on butterflies and the need for their conservation

Across many sub-Saharan African countries, surveys of drivers of butterfly diversity are rare in the literature. In Uganda for instance with a typical wet tropical climate,[24-26,9] showed that there existed no published data describing extensively the diversity of butterflies found in agricultural landscapes in Uganda in relationship to climatic, regional, landscape and local drivers, yet, such information is important for butterfly biodiversity conservation in the rural landscapes. This situation is similar to Zambia where research on butterfly ecology is almost nonexistent. The common literature one is expected to find are field guides such as the publication by [1] a guide to the butterflies of Zambia and to a lesser degree [27] field guide to the butterflies of southern Africa and a handful of other lesser field guides. 

Currently, many studies including those by [28-30] show  that climatic, regional and landscape factors are important drivers for butterflies populations while  other studies also  indicate that local factors (availability of nectaring resources) are of foremost importance in explaining the variation in species richness and total density of butterflies as also earlier noted by[31]       

Although Uganda where the above studies were carried out is wetter than Zambia, the influence of climatic and anthropogenic factors on butterfly populations is the same. What is critical is to isolate and rate or weigh which local (specific local area), landscape, national and regional climatic factors are important predictors of butterfly species richness and abundance in Zambia and particularly the Kafue Flats where this study was carried out. 

1.3 Suitability of butterflies as indicators of climate change

Recent research has shown that butterflies have declined more rapidly than birds and plants emphasizing their potential role as good indicators. Butterflies occur in all habitats in Zambia and so have the potential to act as indicators for a wide range of species and habitats. Unlike most other groups of insects, their taxonomy is understood, they are easy to recognize and we have a wealth of information on their ecology and life-histories. Perhaps this may answer the question, what makes butterflies good indicators? For this reason, we chose butterflies as indicators of climate change. Additionally, because insects in general make up the largest proportion of terrestrial wildlife (more than 50% of species), it is crucial that we assess the fate of insect groups in order to monitor the overall state of biodiversity. Being typical insects, the responses seen in butterflies are more likely to reflect changes amongst other insect groups, and thus the majority of biodiversity, than established indicators such as those based on birds.

 It is also a known fact that insects are particularly useful indicators in the evaluation of landscapes for biological conservation. Among the great diversity of insects, butterflies stand as ideal candidates for ecological study in various landscapes and habitat types. Because of their short life span they quickly respond to environmental variables. For instance, butterflies have been recorded to respond almost instantly to multiple factors such as; plant diversity, habitat complexity, landscape layout, altitude, topography and moisture gradients, and last but not the least climate. 

Protected areas such as National Parks and other land management agencies can use butterflies to evaluate the effectiveness of ecosystems based management. Because of their beauty, when they disappear from the landscape, it is relatively easy to notice their absence.

 The aim of this study was therefore to assess butterfly taxa in relation to local site characteristics and how they influence diversity in different habitats of Lochinvar National Park, as baseline data upon which medium to long-term monitoring of this ubiquitous but important animals group could be based as one way of securing the biodiversity of the Kafue flats. Patterns of butterfly species occurrence and abundance were analyzed to identify habitats and periods of high butterfly activity, diversity, and uniqueness. 

2.1 Study Area Location and Description

2.1.1 Location

       Lochinvar National Park is 410 km² in extent and is geographically located at Latitude 15⁰ 43’ – 16⁰ 01’ South, Longitude 27⁰ 11’ – 27⁰ 19’    East. Altitude, varies between 970 and 1,038 m above sea level (Figure).

Figure 2 Location of Kafue Flats, habitat for the various species of butterflies

2.1.2 Description of the study area

The park is generally flat divided into three major zones, with each zone having a characteristic fauna and flora. The northern third is a flood plain inundated annually by the Kafue River, which also forms the northern boundary. Since 2000, the area has been invaded by alien invasive weed Mimosa pigra.  The central zone comprises termitaria grassland, with grassy plains and scattered termite mounds where Euphorbia candelabrum is common. However, since the construction of the Itezhi tezhi dam, there has been woody encroachment in this zone. The southern zone is a woodland area, which is characterized by mainly species of Acacia, Albizia and Combretum. In this zone, are important and popular geological and historical sites. The Gwisho hot springs are about one kilometer from the park headquarters. The Bwanda group of hot springs lies further west and are said to be among the hottest in Zambia. These hot springs are associated with a deposit of gypsum a substance used in the manufacture of cement. The Gwisho hot springs site was historically occupied by groups of late Stone Age people during the third and second millennia BC, while the Sebanzi hill was a site of a more recent Iron Age village that was inhabited for most of the past 1,000 years. 

About half of the area is part of the Kafue flood plain making a suitable butterfly habitat. Soils are dark grey of alluvial origin. South of the flood plain, is a flat Terminalia zone on sandy clay to clay soils which are water logged during the wet season while hot springs are indicative of a structural fault which occurs near where the woodland meets the southern boundary of the southern edge of the Terminalia zone.  Average annual rainfall is 750mm.

Vegetation is predominantly grass on the flood plain and the most common types are: Oryza birthii, Vossia cuspidata, Echinocloastagnina and Panicum ripens. The commonest herbs are Aeschynomone fluitans and Nymphaea capensis. Steria sphacelata is the characteristic species in the Terminalia grassland. This type of grassland is due to the high water table which is in this zone. South of the National Park is a fire climax woodland of Acacia, Albizia and Combretum spp.

2.1.3 Selection of study sites

This study was carried out on 16 sites inside Lochinvar National Park. The sites selected were; Hand pump, Parks office, baobab, 400m south of Parks  Office, workers compound, Chunga camp, Old lodge, Ecologist's house, main entrance gate, Sebanze hill, airstrip, drum rock, Gwisho road culvert, Chunga lagoon road, Chikolete inside NP, and Munjili area. The sites were located in various habitats representing an array of vegetation communities some affected by various anthropogenic disturbance and management intensities. 

The sites selected were accessible all year round. Established monitoring guidelines for butterfly were employed and no specimen was collected except in very special circumstances where the field guides could not positively aid the identification of a specimen. A Field guide to the butterflies of Zambia and the Field Guide to the Butterflies of [27] were used for identification and nomenclature. Meteorology data for each location were obtained from the National Meteorology Agency in Lusaka.

2.1.4 Data capture techniques 

Field sampling of butterflies was conducted by visiting each of the 16 study sites. Taxonomic characteristics were obtained from the field guides. Surveys were conducted in sunny conditions, usually between 09:00- 16:00hrs when butterflies were more active. Counts were made twice per month at 2-week intervals from January to December.

1. Data Analysis

Descriptive statistics was used to summarize the data and find patterns by obtaining the measures of central tendency such as, mean, median, mode, percentage, frequency and range. 

A non-parametric Chi-square  goodness of fit test was used to compare observed (fo) and expected (fe) frequencies of the identified parameters.  

2.3Ethical Considerations

Specimen collection was only carried out in very exceptional circumstances, particularly when the specimen could not be identified using the field guides. Collection of specimens followed standard protocol. 

3.1 Butterfly Abundance and Diversity 

In total, 96 observations were made in which eight (8) families, 37 genera, and 61 species respectively were recorded (Table 1 & 2).

Table 1:  Butterfly Families recorded in the study area

Table 2: Number of families and their sightings

Lycaenidae, Pieridae and Nymphalidae were the most abundant families and possibly polyphagous in habit, perhaps very well adapted to the environment or may have persisted anthropogenic effects better than others.  On the other hand Hesperiidae butterflies being small, dull colored and very active in flight could be difficult to be observed and may have been missed in some instances, and so were recorded the least. 

3.2 Family frequency observations 

       Of the eight (8) families recorded, the most frequently observed were three; Pieridae, Nymphalidae, and Lycaenidae which had the largest number of genera and species respectively (Figure 1). The differences in the number of observations and number of families were found to be significantly different in favour of the three families (χ² = 37.63, α = 0.05, P < 0.001; χ² = 84.0, α = 0.05, P < 0.001).

Figure 1: Family frequency observation of families, genera and species

3.3 Uniqueness

The three families were unique in terms of frequency of occurrence between sites. Pieridae 23 % (n = 9), Nymaphalidae 20% (n= 8) and Lycaneidae 18% (n=7) giving a cumulative total of 61% of all observations made between sites. The least were; Papilionidae 2.5% (n=1) observed only in one site and the second was Danainae 5% (n=2) observed only on two sites. No family was found to be a generalist as to be common to all sites. 

3.4 Site specific observations

Of the 16 sites visited in this study, three sites accounted for a cumulative total of 51% of species recorded, hand pump 11 % (n = 8), main gate 12% (n = 9) and park office 28% (n = 21). These three sites also had the largest number of genera recorded with a cumulative total of 44% (n = 29), hand pump 6 % (n = 4), main gate 12 % (n = 8) and park office 26% (n = 17). These observations were found to be significantly different (χ² = 74, α = 0.05, P < 0.001) (Table 3)

Table 3: Study site specific observations of families, genera and species

3.5 Comparison across seasons 

Most observations of families, genera and species (70%; n = 67) were made in the dry season between May and September when water sources were limited (Figure 4).  The month of August had the highest number of observation (22%; n = 21) and the lowest were February and March in which no observations were recorded (Figure 4).

Figure 4: Observation across seasons of the year

4.1 Species abundance 

This is the first study to be carried out in this part of Lochinvar National Park, and perhaps the entire Kafue flats area. In that view, therefore, we consider the eight families recorded as a conservatively rich Lepidopteran diversity for a dry savannah habitat such as the Lochinvar National Park which is situated in Agro ecological Region I receiving rainfall not exceeding 700mm per year. [32] in his study of the local habitat characteristics and how they determine butterfly diversity in Kihansi gorge forest of the Udzungwa Mountains in Tanzania recorded only five families with 72 species and considered this as high species richness, although he later acknowledged that the records he obtained in this study were lower than previously published data for this same area in which more than 200 species were recorded. This reduction in species numbers were however attributed to construction and operation of a hydropower facility. [32] he further associated such reduction in species numbers to the 

operations of the power plant and the attendant impacts of environmental change recorded in the gorge ecosystem. Based on these impacts on the lepidopteran taxa, they suggested that the differences in butterfly records was due to the disturbances and temporal changes of conditions in their habitats which may have had influenced decline in abundance of species. They also retaliated that there was no study done to date specifically to quantify the impacts of the reduced river flow on the abundance of food resources such as nectar yielding species and availability of suitable breeding cover such as the presence of larval host plant species., the impacts of which have been observed and recorded in other areas as earlier documented by Kitahara, [33]

4.1.2 Importance of water availability 

In this study, a preponderance of skewness towards sites with permanent water all year round was observed. There were more and frequent observations in sites with water availability throughout the year than sites without water in the dry season. This is because butterflies have an affinity for water, because they need to take either food or water for their liquid diet. Anthropogenic modifications to the environment such as the provision of water taps played a role in determining butterfly populations distribution and frequency of occurrence. 

This explains why most observations were recorded at the Park offices which have water taps that more often than not leak providing a permanent water source and plants growing nearby often being green throughout the year. The hand pump is also a site with a permeant source of water as staff and their families draw water from the hand pump and the overflow often creates a permanent pool of water which the butterflies access. A similar situation obtains at the main gate. The other sites depend on rain water and often remain dry for a large part of the year. 

4.1.3 Seasonal Population Dynamics

Seasonal population dynamics was observed as butterfly numbers were more in May-September but less from December to March. The decline in butterfly abundance from December to March may be attributed to the life cycle.

Butterfly and moth taxa have a four (4) life cycle or stages, egg, larva (or caterpillar), pupa and adult. This cycle is called complete metamorphosis. It is likely therefore, that at the beginning and during the rainy season most species would be in their egg, or larva stages, and then towards the end of the rainy season they would be in pupal stage. This may account for the low to no observations in some months of the year as was observed during this study. In this study, the months of February and March had no observations. It is very likely that most genera were in caterpillar stage as this is the time they feed on green vegetable matter and this is also the time that some species cause damage to crops as pests. 

genera and species were expected to be recorded as is the case with several other studies in other areas where fewer families were recorded but with a large number of species. Perhaps, savannah conditions of dryness and less cover could be a plausible reason for this, because forested areas often host more butterfly diversity and populations. This assumption is supported by the fact that butterflies are primarily a tropical group of insects and often depend on rain forest plants for larval food. Similarly, according to studies from Manembo - Nembo Wildlife Reserve (Indonesia), more abundant and diversified butterflies were found in riverside habitats in the forest, which is a source of food and provides shelter. In contrast, in Sicily, Italy, grasslands were found to be the most suitable for butterflies because grasslands provide the food and shelter the larvae need. 

In this study, the common habitats recorded were open grassland, residential areas, and some bushland and thicket communities which may practically not support high butterfly density compared with forested areas which support butterflies more effectively because of the variety of biomass they provide.

Ecologically, insect species diversity correlates strongly with the structural complexity of habitats and the diversity of vegetation. Food quality also may influences their occurrence. So the, high butterfly diversity reported in forest ecosystems is in response to food quality, as the study by [34] showed that butterflies were not found in disturbed lands; which also supports the idea that degraded habitats affect the population of butterflies. Generally, the diversity of butterflies depends on habitat management and butterfly species richness increases with increasing diversity of plants and decreases with increasing degradation of vegetation.

In this research work therefore, we conclude by suggesting that the results of this study in which we treated the eight families recorded as high diversity should be noted with great care, because, the area has experienced tremendous anthropogenic impacts over the years such as housing, road network construction and others as well as the habitat transformation of the entire Kafue flats after the construction and commissioning of the Itezhi-tezhi hydroelectricity dam in 1979. 

The other contemporary factor we should consider is the impact of climate change and its effect on habitat quality in the last few decades. Climate change effects can operate directly through changes in aspects of climate, particularly those that are heat-related, and through changes in habitat diversity. Earlier conceptual models by [35] suggest that regional-scale habitat diversity will decline in a rapidly warming climate and therefore, our remarks on diversity and species richness can only be confirmed with continued monitoring in the future and the continuous review of the isolated records in the past. By reviewing previous records including those relating to temperature rise, it may be possible to estimate previous losses or reduction of some taxa, because physiologically, butterflies and their caterpillars which is the second phase in the process of metamorphosis have an optimum temperature range, within which body processes function optimally. If there are significant local microclimate changes, it may affect their survival and thus have an effect on their numbers and spatial distribution.

       We also take note of the influence of the abiotic environment: For instance, drought-periods as well as heavy showers and flooding as has been experienced in the last three years (2019 -2022) in southern Africa affecting mainly Mozambique, Malawi and to a lesser degree Zambia, could alter habitat factors suitable for butterflies. The direct impact would be vegetation structure as it relates to, in some instances nectar yielding plant species which would affect foraging. Therefore, the changing rainfall patterns, increased frequency of droughts we have experienced in the recent past may have had an influence or altered species composition and spatial distribution of some plants and therefore lepidopterans depending on them. If this did occur, then one would assume a low diversity as being the ultimate result. 

4.3 Recommendations

In making recommendations for future monitoring and management of lepidopterans, we emphasise the need for the assessment of local drivers influencing richness and abundance of butterflies such as the abundance and distribution of floral nectaring resources among others. Future studies therefore should consider assessing percent cover of flowering plants (trees, shrubs, herbs and weeds), and the number of nectaring plant species. In areas outside the PA it may be worthwhile considering the percentage cover of cultivated floral resources (proportion of cultivated pollinator-dependent and non-pollinator-dependent crops) per unit area.

As regards climate change, cumulative evidence suggests major effects on lepidopterans, leading to species declines and extinctions in some instances as earlier mentioned by [36,23] predicted that the impact of the expected future changes on the lepidopteran populations and distribution will be huge, and action was urgently needed to address this concern. Therefore, measuring the effect of climate change should constitute the first critical step in understanding how butterflies are affected, the speed at which such changes are taking place and the time needed to take amelioration measures.  To actualize this on the ground, [37] suggested several methods that could be utilized to study the effect of a changing temperature on lepidopterans. Worth exploring is the method of [38] in which the change of the Community Temperature Index (CTI) proved to be the most sensitive and easy to use indicator for the effect of climate change on butterflies. This is one possibility the management staff of Lochinvar National Park may consider using among other tools. 

The park staff should initiate collaboration with butterfly enthusiasts and Nature Based NGOs to work towards promoting awareness among the populations so that land barriers that may hinder mobility across landscapes are minimized and land use practices that mimic agri-environments promoted. The awareness campaigns’ should be widened out to underscore the concern that since lepidopterans are good biological indicators, holistic measures to help them will help conserve Zambia’s rich biodiversity. 

Enhance preservation of the natural habitats such as the Lochinvar National Park, which offers a large and diverse set of habitats with a mosaic of microclimatic conditions, can undoubtedly help sustain larger and more stable butterfly populations and communities for a much longer time than small areas particularly in isolated urban habitats. This would also perhaps help facilitate connectivity between areas with large butterfly populations such as protected areas, private land and land under customary tenure where lepidopterans occur in substantial numbers. Greening urban areas where this is feasible should be encourage as this is also likely to achieve similar results.

Away from mega fauna, local scientists should be enticed to supplement the efforts of entomologists who are limited in number to carry out research on butterflies which would in turn provide a frame work to monitor their ecology and to follow changes arising from the impact of climate change and also as a way of building a robust and representative set of indicators. 

This baseline study, it is suggested, should be extended to other protected areas so as to further understand the impact and response of lepidopterans to the current anthropogenic activities and climate change.

Author contributions

Both authors contributed equally to conceiving the study, collecting and interpreting data, and developing the final manuscript. 

Acknowledgements

We acknowledge the contributions of field staff at Lochnivar National Park who toiled in visiting the 16 sites and many others who critiqued the document from its inception to the final stages. 

The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest. This was a self-funded research activity. 

No funding sources

The study was approved by the

Mulungushi university.

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