The Dragonflies and Damselflies of Angola: An Updated Synthesis

Prior to 2012, only 158 species of Odonata were known from Angola. Surveys in 2012 and 2013 added 76 species and further additions in 2016 brought the national total to 236 species. This was published earlier in 2017 as the checklist of the dragonflies and damselflies (Odonata) of Angola by the same authors (Kipping et al. Afr Invertebr 58 (I):65–91. https://africaninvertebrates.pensoft.net/article/11382/, 2017) on which this chapter is based. Records obtained in 2017 and 2018 and a survey by two of the authors in December 2017 led to the discovery of 25 additional species, of which several are undescribed. We provide a revised checklist here comprising 260 species and discuss the history of research, the biogeography of the fauna with endemism and the potential for further discoveries. The national total is likely to be above 300 species. This would make Angola one of the richest countries for Odonata in Africa.


Introduction
Given the country's size, diverse landscapes, climatic regimes and habitats Angola is likely to be one of the richest in Odonata species in Africa. However, Angola's biodiversity is very poorly known, with comparatively limited research before independence in 1975 halting altogether in the three decades of civil war and unrest that followed. Research coverage is also limited for Odonata, with much of the north and east never surveyed at all (Clausnitzer et al. 2012). The potentially very species-rich highland catchments of the Congo, Cuanza, Cubango (Okavango) and Zambezi rivers are almost unknown and may hold many undescribed species. The whole Angolan part of the extensively marshy Cuando River and almost the entire Cuito River system are also largely unsurveyed.

History of Odonata Research in Angola
Research on Odonata began in July 1928, when the Swiss zoologist Albert Monard embarked on the first of his two expeditions to Angola, which lasted until February 1929. Monard was a curator at the Natural Museum of La-Chaux-de-Fonds in Switzerland with a broad interest in nature who mainly collected vertebrates and plants. Ris (1931) identified 27 and described four species from Monard's first expedition.
With the death of Ris, Monard submitted the Odonata from his second expedition (April 1932to October 1933 to Cynthia Longfield at the British Museum (now the Natural History Museum) in London, who had published several records obtained by Karl Jordan from Mount Moco in 1934 (Longfield 1936). Longfield (1947) identified 77 species from Monard's new material and described 13 new species and two new genera. She also dealt with the Odonata held at the Dundo Museum in northern Angola, first revising the genus Orthetrum based on the long series available (Longfield 1955) and later listing 61 species from the collection, including three new species (Longfield 1959).
Elliott CG Pinhey (1961a, b) described five new species of Gomphidae from northern Angola received from António de Barros Machado of the Dundo Museum. While Longfield (1959) stated that the Dundo collection "shows the usual scarcity of the genera Gomphidae", Pinhey (1961a) noted it "was particularly notable for the number of Gomphids." Possibly Machado split the material between the two authors. It is uncertain whether the material was collected in Dundo or only held there, as most records lack details on collector, date and precise locality. However, Pinhey (1961b) did detail collecting in localities around Dundo, suggesting that all material came from this part of Lunda-Norte Province. The collector was probably Machado himself. No-one has worked on this collection since and its state is thus unknown. Elliot Pinhey was curator at the National Museum of Zimbabwe from 1955 until 1975 and while he collected intensively in adjacent countries, he only visited Angola twice (Vick et al. 2001). In April and May 1963 Pinhey participated in an expedition to northwestern Zambia, also visiting an area east of Caianda and the Lutchigena River in Angola directly adjacent to the Ikelenge Pedicle of Zambia, where he recorded 26 species (Pinhey 1964(Pinhey , 1974(Pinhey , 1984. His second excursion into Angola went to an area between Luanda and the Duque de Bragança Falls on the Lucala River (now known as Calandula Falls) in October 1964 with records of 32 species (Pinhey 1965).
Pinhey further treated the material of three collectors, describing a species in honour of each of them. Edward S Ross of the California Academy of Sciences collected between Cuchi and Dondo in 1957and 1958(Pinhey 1966, the American expert of mammal behaviour Richard Estes in central Angola in 1970 (Pinhey 1971a), and Ivan Bampton around the Serra da Chela and Tundavala in 1973 (Pinhey 1975). In the 1975 paper he also repeated records from Pinhey (1964Pinhey ( , 1965 and Longfield (1947), and provided a gazetteer, causing confusion about the precise locality of some sites. The correct historic collecting sites could be verified with the gazetteer of Mendes et al. (2013).
Various collectors gathered about 1000 specimens in the collection of the Instituto de Investigação Agronómica in Huambo between 1950 and 1974. These records were never published but this will be done shortly by Sara F Elizalde and David Elizalde as a GBIF dataset.
After Angola's independence in 1975 there was a long break in field research, with only a few records by various collectors. Namely in the two decades between 1980 and 2000 not a single record of Odonata is available. Some years after the end of the civil war a renaissance of research began, resulting in a growing number of records ( Fig. 9.1). All localities with available Odonata records distinguished in the three periods (a) pre-independence 1928-1974, (b) after independence 1975-2001 and (c) after the end of civil war 2002-today are shown in Fig. 9.2.

Origin of Recent Data
In January 2009, an expedition led by Brian Huntley visited the Serra da Chela in southwestern Angola and the Namib Desert to the south. During that survey Warwick Tarboton collected and photographed Odonata around Humpata (7 field days).
Jens Kipping surveyed the upper catchment of the Okavango (Cubango) River on the SAREP (Southern African Regional Environmental Program) Expedition from 5 to 22 May 2012 (18 field days). A second SAREP survey visited southeastern Angola with the Cubango and Cuando River floodplains in April 2013.  The National Geographic Okavango Wilderness Project (NGOWP 2018) gathered scattered data on Odonata which has not yet been fully considered, except for some field photographs and exceptional records provided by John Mendelsohn.
Apart from the field surveys the authors also examined the Angolan collections and type material in the Natural History Museum in London, the National History Museum of Zimbabwe in Bulawayo (Dijkstra 2007a, b), the Royal Museum for Central Africa in Tervuren, Belgium and the Instituto de Investigação Agronómica in Huambo, Angola. All records are kept in the Odonata Database of Africa -ODA (Kipping et al. 2009) and mapped per species on African Dragonflies and Damselflies Online -ADDO (visit http://addo.adu.org.za/ also for further information about all mentioned species).

Odonata Species Recorded in Angola
From all the historic sources mentioned above, 152 species of Odonata were known to occur in Angola until 2009. Some of the formerly published species had to be deleted from the country list in the light of new taxonomic knowledge and after careful validation of all records (see Kipping et al. 2017).
In 2009 Warwick Tarboton recorded 47 species of Odonata at the Serra da Chela of which five were recorded in Angola for the first time and one was new to science (Tarboton 2009, Dijkstra et al. 2015. The first SAREP Expedition in 2012 yielded 87 species, 17 of them new to the country list and two new to science (Kipping 2012, Dijkstra et al. 2015. One additional species new for the country came from a second SAREP Expedition in April 2013 of which all collected specimens were examined. The first expedition to Uíge, Negage and Ndalatando resulted in 138 species, of which 43 were recorded for the first time in Angola and five were new to science. The second visit produced 86 species, adding another 15 to the national list. The updated checklist of the Odonata of Angola, now of 260 species, is provided in Appendix 1. The ODA database now holds about 4900 Angolan records from more than 400 localities. All species of Appendix 1 are reliably recorded from Angolan territory. Footnotes will give further information about the 25 additional species to the updated country list and one species that was deleted from it. There are 15 more species listed in Appendix 2 that are known to occur at rivers bordering the country with Namibia and Zambia. The Namibian bank of the Okavango River is very well surveyed Martens 2007, 2014) and most of the mentioned species derive from this river. These species were technically not found on the Angolan riverbank and therefore not included in the checklist. But naturally they belong to the country's fauna.

Composition
Angola's rich dragonfly fauna expresses its geographic position, size and diversity. Its territory, especially in the north, falls within a region with an estimated highly diverse fauna ( Fig. 9.3). Dijkstra et al. (2011) observed that roughly half of tropical African species occur predominantly within the extensive lowland forests of western and central Africa, a quarter is associated with the eastern and southern part dominated by highlands, while the remaining quarter occurs in open habitats throughout much of the Afrotropics. Indeed, about half of Angola's species are widespread across the continent and its exceptional diversity can be attributed to two major sources. Almost 30% are confined to forest habitats in the north, mostly below 1000 m altitude. Nine species confined to the Lower Guinea, the forest area that stretches between the Congo Basin and Atlantic Ocean from Cameroon to Gabon and western Congo, reaching their southern limit in northwestern Angola. Nearly 20% favour the swamps, grasslands, miombo woodlands and gallery forests that stretch eastwards, mostly above 1000 m asl. This fauna is concentrated in Katanga and northern Zambia but has now been proven to extend across to the Angolan upland. This was confirmed by the discovery of Orthetrum kafwi at two localities in Cuanza-Sul in December 2017. The species was until then only known from Upemba NP in southern DRC. A number of palustrine species, e.g. Anax bangweuluensis, Pinheyagrion angolicum, Pseudagrion deningi and P. rufostigma, prefer larger marsh areas and swamps as in the Okavango Delta of Botswana (see Kipping 2010) and spreading north into Bié highlands with the headwaters of that river system. The discovery of Trithemis integra near Uíge is also of special interest, as it had seemed to be endemic to the Albertine Rift, being known previously only from western Tanzania and Uganda and eastern DRC.

Endemism
Seventeen valid species and several recently discovered undescribed species have so far only been found in Angola (Figs. 10.4,10. 5 and 10.6 for examples). With the exception of two known only from their alleged type localities in far northeastern Angola, all are limited to the central plateau: the type locality for Platycypha rubriventris is questionable as it may be that of Pseudagrion dundoense, which could also be a river species from the very poorly sampled southern Congo Basin. No endemics have been found below 1200 m asl in the east, although some drop down to about 500 m west of the escarpment. While the proportion of endemics (7%) is lower than for Ethiopia (12 endemics; 11%) and South Africa (30 endemics; 18%), countries that also enclose distinct highland areas, this still ranks Angola as one of Africa's greatest centres of endemism for Odonata, rivalling the highlands of Cameroon (13 endemics) and the Albertine Rift, Eastern Arc and Katanga. Moreover, the number is expected to increase, as almost two-fifths were described since exploration was reinitiated and undescribed species of Platycypha, Paragomphus and Tetrathemis are already known to us.
Only Platycypha presents an endemic radiation. While Chlorocypha (the family's other large Afrotropical genus) has diversified with almost 30 species largely in the forested lowlands of west and central Africa, Platycypha is ecologically more diverse, with species adapted to open, submontane and lake habitats as well. The Angolan endemics are found mainly between 1300 and 1800 m altitude in open habitats. The widespread P. angolensis replaces the common P. caligata, which extends from South Africa to Ethiopia but only peripherally into Angola. Platycypha bamptoni is probably confined to Serra da Chela; a similar undescribed species appears more widespread. Platycypha crocea is typical of very small streams in the Bié highlands and escarpment mountains whereas the other two inhabit larger streams and rivers. A local radiation of a group that has otherwise diversified in the highlands to the east, and forests to the north, fits the overall affinities of Angola's endemic Odonata both geographically and ecologically (Kipping et al. 2017) (Fig. 9.4).
The four endemic Pseudagrion species have separate origins but similar links: the nearest relatives of P. angolense and P. estesi appear to be the rainforest species P. grilloti Legrand, 1987 and P. kibalense respectively. The former is limited to Congo and Gabon, but the latter extends to Cameroon and Uganda. P. sarepi is closely related to P. fisheri and P. greeni, both of which extend from Angola into Zambia. While these species belong to the genus's A-group, the B-group species P. dundoense is known only from Dundo and may not be endemic at all (see above).
Notogomphus kimpavita is the sister-species of N. praetorius found in highlands across southern Africa (including Angola), while Eleuthemis eogaster is nearest to an unnamed species from Gabon (Dijkstra et al. 2015).
Molecular data for Umma femina and Onychogomphus rossii are not available yet. By its unusual habitus and colouration Umma femina (see Fig. 9.5) is a very distinct member of the genus. It is definitely the odonate flagship species of the Angolan highlands and probably also most threatened. The morphology of O. rossii is close to other pale Onychogomphus species from the open plateaus stretching from Angola to Zambia and Katanga.
Thus, like the majority of Angola's Odonata, most endemics probably originated quite recently and proximally from the forests to the north and open habitats to the east. However, some affinities are unresolved and potentially more distant: Agriocnemis toto and especially A. canuango have no obvious close relatives (Dijkstra et al. 2015), while the near-endemic A. angolensis and A. bumhilli are probably related to each other but even more distinct overall (Kipping et al. 2017). These data suggest that Angola may be the centre of diversification of this genus, which includes Africa's smallest damselflies. The morphologically very distinct Aciagrion rarum is only known from very few specimens from Lunda Sul Province in the northeast and molecular data is not available yet.
Micromacromia flava is morphologically nearest M. miraculosa (Förster 1906), known only from the East Usambara Mountains of north-eastern Tanzania and the only one of four Micromacromia species adapted to non-forest habitats, being strongly pruinose with maturity. Elattoneura tarbotonorum may be closest to E. frenulata of southwestern South Africa (Dijkstra et al. 2015): after its discovery at the Serra da Chela in 2009 it was found more widespread in December 2017 along the mountain range stretching north into Cuanza-Sul.

Potential for Discovery
If we compare the tallies for the well-studied neighbouring countries of Zambia and Namibia, the total number of species in Angola should lie somewhat above 300, meaning that less than 80% of the fauna is currently known. All Odonata expeditions in modern times surveyed areas that are easily accessible.
Additions can be expected throughout the country, but especially in the remote regions on the eastern and particularly northern border, as species diversity is expected to be extraordinarily high in the transition to the Congolian rainforest ( Fig. 9.3). The province of Lunda-Norte with only 92 recorded species and 162 records should be the richest area for discovery, around Dundo where exploration began in the 1950s. Generally, all the northern and eastern provinces are largely unsurveyed and the discrepancy between the amount of available data, the number of known species and the expected diversity is extremely high. This applies also to the provinces of Lunda-Sul (10 species, 11 records), Zaire (17 species, 21 records), Malanje (35 species, 152 records) and Moxico (46 species, 51 records). An exception is Uíge where recent surveys increased the number of known species to 145 from 820 records gathered.
The central highlands can also yield more surprises, like the discovery of additional endemic species, with three areas being especially notable. Firstly, despite having most records, the north-south directed mountain range that lies entirely above 1600 m asl and includes the Serra do Chilengue, Serra da Chela and Angola's highest peak at Mount Moco (2620 m asl) is poorly sampled as the large gaps in Fig. 9.2 illustrates.
Secondly, except for its extreme northern and southern ends, the western escarpment has only been surveyed recently, which already led to the discovery of an undescribed Paragomphus species from Cumbira Forest. Even more easily accessible provinces such as Bengo and Cuanza-Sul will prove to be much richer in species than currently known. The potential of these mountains is illustrated by the discovery of a spectacular and unique but unknown species by Chris Hines and Rogério Ferreira in May 2018 ( Fig. 9.6). Two males at a stream that flows off the Namba Mountains in Cuanza-Sul were photographed but not collected. These mountains reach over 2000 m in altitude and harbour larger pockets of Afromontane forest than Mt. Moco (Mills et al. 2013). They are known for their plant endemism and fieldwork there will definitely lead to the discovery of more endemic Odonata.
Thirdly, an extensive plateau at 1200-1600 m altitude stretches east from the Bié Highlands. Except for its southern edge, this area shared between Bié and Moxico Provinces, which is almost as large as Uganda (or the United Kingdom), has almost no records. A few collections from the NGS Okavango Wilderness Project suggest that more new species for the country and for science can be expected here. These deep Kalahari sands are the 'watertower' of Angola and its neighbours, incorporating the headwaters of the Cuito, Cuando, Chicapa, Cuango, Cuanza and large tributaries of the Congo and Zambezi such as the Cassai and Lungué-Bungo. The sources of the vast catchments of the Congo, Cuanza, Okavango and Zambezi meet in a small area between Munhango and Cangonga. Watersheds are prone to endemism (Dijkstra et al. 2011) and this region is the top priority for further research.
Studying insect collections of Angola's museums will be also a valuable source of more records and possibly even to get insight into past conditions in the light of the recent landscape change. Of special interest is the Dundo Museum that holds many interesting specimens of Odonata, some of which have been published. This remarkable collection has not been studied since independence but survived the civil war. There is also material dispersed over several museums in Europe and probably also in private collections, mainly in Portugal.
New species are most likely to be found among genera prone to narrow (highland) ranges, i.e. with known Angolan endemics like Platycypha and Pseudagrion, but also Agriocnemis, Elattoneura, Notogomphus and Paragomphus. Also possible Fig. 9.6 An undescribed species that probably belongs to Trithemis, although the extensive markings and dense veins in the wings are unusual even for that highly diverse genus. Two males were observed at a stream running off Namba Mountains in Cuanza-Sul. (Photograph by R Ferreira) is the discovery of endemics in genera that are well represented across the country and continent, and that have highland endemics elsewhere but not in Angola, such as Africallagma, Neodythemis and Orthetrum. However, given the biogeographic diversity of Angola's fauna and endemics, we could expect greater surprises. Among forest genera with no known Angolan endemic, Allocnemis seems most likely to reveal one, e.g. on the escarpment. The presence (or local endemism) of distinctly Lower Guinean genera like Neurolestes, Africocypha, Pentaphlebia and Stenocnemis seems less likely, but the Lower Guinean Stenocypha gracilis (Karsch 1899) has four endemic relatives in the Albertine Rift and the sister-taxon of the Upper and Lower Guinean Tragogomphus is Nepogomphoides stuhlmanni (Karsch 1899) in the Eastern Arc, suggesting an Angolan taxon is possible.
Some typical African highland genera are notably absent from Angola. Atoconeura is most likely to be present above 1400 m asl, being found in Zambia, Katanga, the Lower Guinea and Albertine Rift. However, its absence also from South Africa suggests historical factors may have been limiting, e.g. that the highlands were too harsh in cooler periods and too isolated when habitats were suitable (Dijkstra 2006).
This might not apply to Proischnura, present in South Africa as well as Cameroon and the Albertine Rift. However, that genus is absent from Katanga and northern Zambia, which lies lower and thus possibly provided no stepping-stone to the mountains of Angola. Kipping et al. (2017)

Conservation
Our findings show that Angola's wealth of aquatic habitats harbours a rich freshwater fauna. Although large areas are relatively untouched, Angola's rapid economic and population growth will have a tremendous impact on the environment and thus human well-being in the future. In the light of this, Angola's development should consider (1) the establishment of sewage works in cities and larger villages; (2) a stop to deforestation, especially along stream courses; (3) restoration of deforested water catchments; (4) village-level awareness campaigns for sustainable use of freshwater sources, e.g. no detergents and waste dumping in rivers; (5) biodiversity surveys and monitoring to feed into a national conservation plan.
With the exception of four species, all endemics are currently considered Data Deficient for the IUCN Red List of Threatened Species. Platycypha angolensis, Pseudagrion angolense and Micromacromia flava are Near Threatened because, while they seem fairly widespread, their dependence on relatively natural habitats may put them at risk as human development progresses. Only Umma femina is now listed as threatened. It is currently known from only a few sites in the fairly densely populated highlands around Lubango and seems to inhabit exclusively the smaller and cooler highland streams. There is much development in this densely settled region and increasing pressure on those habitats by grazing, deforestation and urbanisation. As it seems to prefer cool mountain streams we can assume additional risks from climate change and it is therefore thought to be Vulnerable to extinction. More research on all endemic species' statuses and ecology is urgently required.
Angola has an exceptional fauna of dragonflies and damselflies, as well as many valuable rivers and wetlands. Odonata are excellent indicators of the health and biodiversity of both the freshwater and terrestrial realm. As the biological survey of Angola advances, they should be a priority taxon. Rural Development and Fisheries) for their help in organization and management on the 2012 tour, and to Vince Shacks and Werner Conradie for collecting specimens on the second SAREP survey in 2013. We thank Alvaro Bruno Toto Nienguesso, the driving force behind biodiversity research in Uíge Province, Angola, Prof Dr. Neinhuis and Dr. Thea Lautenschlaeger from TU Dresden for inviting us to the field survey in Uíge province. Part of the fieldwork in Angola was supported by a travel fund from the German Academic Exchange Service (DAAD). We thank Dr. Aristófanes Pontes, director of the Instituto Nacional da Biodiversidade e Áreas de Conservação (INBAC) for the support during the December 2017 expedition, by providing the necessary permits. Chris Hines provided many valuable photographic records. Further records were provided by Warwick Tarboton, Dr. Manfred Haacks (SASSCAL), John Mendelsohn (RAISON) and Rogério Ferreira. The latter also gave permission to use his wonderful photograph.

Appendix 1
Checklist of Odonata recorded from Angola.
# -see Taxonomic comments in Kipping et al. (2017); ## -see footnotes following this table; (V) Validation of species: "1!" new national record made by the authors; "1!!" new national record made by the authors and addendum to Kipping et al. (2017); "1" records obtained by authors and confirming existing records; "2" specimens kept in collections (identification confirmed or primary types); "3" literature records, regarded as reliable because specimens were described well or location agrees with known biogeographic pattern; "4!!" new national record made by other persons and addendum to Kipping et al. (2017); ** -range restricted to Angola; * -range restricted to Angola with very few exceptions (see Endemism in the discussion).
Zosteraeschna minuscula -male collected at the Tchiamena River near Lubango on Serra da Chela, Huíla Province in December 2017. Northernmost record; widespread in South Africa but with scattered records in Namibia and eastern Botswana.
Paragomphus cognatus -presence in Angola was uncertain due to lack of reliable material (Kipping et al. 2017), but several males collected at Tchiamena, Leba and Neve Rivers in the Serra da Chela in December 2017.
Paragomphus sp. nov. -two males collected at the Uiri River near Conda, Cuanza-Sul Province in December 2017 belong to an undescribed species similar to P. cognatus but darker and with stouter paraprocts and more curved cerci.
Bradinopyga strachani -the Instituto de Investigação Agronómica in Huambo has three males from Ndalatando, Cuanza-Norte Province from March 1973, coll. U Passos. Several also collected at Rio Mussenju, south of Quilengues, Benguela Province in December 2017 and photographed by R Ferreira at Calandula Falls, Lunda-Norte Province in June 2018.
Orthetrum kafwi -several males and females collected at boggy streams and bogs in the highlands around Cassongue, Cuanza-Sul Province in December 2017. Previously know only from the type locality in the Upemba National Park in Katanga, which lies 1400 km to the east.
Rhyothemis cf. notata -male photographed by J Mendelsohn at Sacangombe near the Cuito River source in Moxico Province in November 2011. The black markings in the forewings reach only to the nodus and in the hindwings halfway the nodus and pterostigma, which is much less than even the palest variation of R. notata illustrated by Dijkstra & Clausnitzer (2014). The habitat is open, while true R. notata favour rainforest conditions. This species therefore needs to be verified with specimens.
Tetrathemis sp. nov. -several males photographed by C Hines in dry forest near Cambondo, Cuanza-Norte Province in March 2017. Differs from T. fraseri by the smoky wings and shape of the very hairy cerci.
Trithemis aenea -photographed by C Hines near Lucala north of Uíge in June 2017.

Appendix 2
Odonata recorded from rivers bordering Angola that most likely also occur in Angola.
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