Research Article |
Corresponding author: Rafał Bernard ( rbernard@amu.edu.pl ) Academic editor: Pavel Stoev
© 2018 Rafał Bernard, Bogusław Daraż.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Bernard R, Daraż B (2018) New records of dragonflies (Odonata) in Zambia. African Invertebrates 59(2): 165-193. https://doi.org/10.3897/afrinvertebr.59.29021
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Zoogeographically important data on the occurrence of 22 dragonfly species in Zambia are presented, including at least seven species for the first time recorded or unambiguously confirmed in the country. They filled gaps in the previously known distribution ranges and showed that some of them reach further, especially to the south, but also west or north. Zoogeographical considerations are completed with some remarks on species’ morphological traits and habitat selection and activity.
Africa, Afrotropical fauna, zoogeography, Zygoptera , Anisoptera , Gynacanthini
Studies of Afrotropical odonates have been significantly intensified since the end of the 20th century. Apart from many taxonomic works, they brought the spectacular event of publication on 60 new species for science (
In contrast to adjacent countries, the published knowledge of odonate fauna of Zambia has still remained on a relatively low level. It is still mostly based on the synthetic paper from the 1980s (
We organised three expeditions to central Zambia in various phases of the rainy season, in December 2013, April 2015 and January 2017. Apart from several localities visited on the way (in and near Kabwe), our studies were intentionally focused on a small remote area combining the bottom of the rift valley (the Luano Valley), its mountain slopes (the Muchinga Escarpment) and the adjacent upland plateau. In total, 107 species were recorded, but the whole material will be described in a separate paper analysing the species occurrence with reference to habitats in a small territorial scale. In the present paper, 22 selected species are presented, whose newly discovered localities are zoogeographically important to better explain the species distribution and to fill gaps or expand limits in the known pictures of ranges. Zoogeographical considerations are completed with some remarks on the morphological traits and habitat selection and activity of species.
All the investigated localities are situated in the Central Province of Zambia and almost all, apart from the last (No. 27 for Trithemis nuptialis), in broad environs of Chingombe, Luano District. Numbers of localities are given in square brackets.
Collected males are abbreviated to M and females to F. These individuals are in the collection of Rafał Bernard in the Nature Collections of the Faculty of Biology, Adam Mickiewicz University in Poznań, Poland.
An entry “1 M, 1 F” means that one male and one female were observed and collected. Another kind of entry, “3 ovipositing females (1 F)” means that three females were observed and one of them was collected.
Dimensions are given in millimetres. Abdominal length excludes appendages. The following abbreviations are used: Abd = abdominal length; S1–10 = abdominal segments 1–10; Fw = forewing; Hw = hindwing (when used with a number means hindwing length); Pt = pterostigma; Ax = antenodal cross-veins; Mx = median cross-veins; Al = anal loop cells; At = anal triangle cells; if the number of Ax, Mx, Al and At does not differ between the left and right wing of an individual, it is given as one number, (e.g. 31), but if it differs between wings of an individual, it is given for each wing in a pair separately, (e.g. 21/22). BMNH = The Natural History Museum, London, UK; MRAC = Musée royal de l’Afrique centrale, Tervuren, Belgium.
COI sequences of Notogomphus cf. zernyi and Gynacantha sp. were obtained with the use of standard procedures and compared with the known (partly unpublished) sequences from these genera in BOLD: the Barcode of Life Data System (
Data on the distribution of Odonata in Africa were taken from the Odonata Database of Africa (ODA) illustrated on maps of the ADDO website.
[1.] Dambo Kabufumu, 34.5 km WNW of the Chingombe catholic mission, by the road to Old Mkushi; 14°23.31'S, 29°38.96'E, 1,230 m a.s.l. – the plateau; a large (90×30 m) temporary pool in the dambo, i.e. in a river-like open valley, partly inundated with water in the wet season, between slopes with miombo woodlands. With crystal water up to 1 m deep and with abundant Cyperus sp., grasses and a large patch of Persicaria senegalensis f. albotomentosa.
[2.] Dambo Kabasa, 28.5 km WNW of the Chingombe catholic mission, by the road to Old Mkushi; 14°23.04'S, 29°42.23'E, 1,200 m a.s.l. – the plateau; a large (50×45 m) pool in the dambo. In 2015 and preceding years, highly astatic but probably semi-permanent. With rich and abundant vegetation: emergent grassy and Schoenoplectus-like along shores, nymphaeids Nymphaea caerulea on the water table and submerged Ottelia ulvifolia.
[3.] The Mikwa Stream, uppermost reach, just below a source, 8.3 km W of the Chingombe catholic mission; 14°25.27'S, 29°53.39'E (to 100 m upstream and downstream), 1,180 m a.s.l. – the mountains. Permanent, narrow (0.5–2 m), with heterogeneous morphology (boulders, cascades, strong-current sections and calm deeper stream pools) and correspondingly diverse bottom deposits from gravelly and sandy up to muddy with a thick layer of detritus. Locally with side seepages. Deeply shaded by a gallery forest, with small clearings.
[4.] The Mikwa Stream, upper reach, 8.0 km W of the Chingombe catholic mission; 14°25.15'S, 29°53.51'E, 1,160 m a.s.l. – the mountains. For the habitat description see [3.].
[5.] The Mikwa Stream, upper reach, 7.7 km W of the Chingombe catholic mission; 14°25.04'S, 29°53.67'E, 1,120 m a.s.l. – the mountains. For the habitat description see [3.]. Additionally, the main stream fed by a smaller watercourse and side trickles.
[6.] A small tributary of the Mikwa Stream, 7.5 km W of the Chingombe catholic mission; 14°25.03'S, 29°53.81'E, 1,110 m a.s.l. – the mountains. The narrow (0.3–0.6 m, exceptionally 1.5 m) headwater mountain stream: steep, with cascades and shelves, calmer and faster sections, sandy bottom, locally with detritus. Shaded with clearings.
[7.] A pool situated next to a small tributary of the Mikwa Stream, 7.5 km W of the Chingombe catholic mission; 14°25.03'S, 29°53.81'E, 1,110 m a.s.l. – the mountains. Small (1.8 m2), temporarily fed by a trickle. With clear water (30–40 cm deep) and a lot of old leaves. Shaded with clearings.
[8.] Seepage feeding a tributary of the Mikwa Stream, 4.6 km W of the Chingombe catholic mission; 14°25.22'S, 29°55.40'E, 680 m a.s.l. – the mountains. Small (7 m2), flowing from under a rock, with warm water and loose vegetation. Partly sunny.
[9.] The Mikwa Stream, lower reach, 4.6 km W of the Chingombe catholic mission; 14°25.23'S, 29°55.45'E, 670 m a.s.l. – the mountains. A few metres broad, with gravelly-stony bottom, boulders and rapids. Shaded by a gallery forest.
[10.] A pool ‘hanging’ in an earth-slip on the slope of the Mikwa Stream valley, 10 m above the stream, 4.2 km W of the Chingombe catholic mission; 14°25.17'S, 29°55.63'E, 640 m a.s.l. – the mountains. Small (4–5 m2), deep (>1 m). Water not transparent, warm, fed by a warm-water trickle. Shaded with clearings.
[11.] A seepage trickle (feeding a small deep pool) on the slope of the Mikwa Stream valley, 10 m above the stream level, 4.2 km W of the Chingombe catholic mission; 14°25.17'S, 29°55.63'E, 640 m a.s.l. – the mountains.
[12.] A complex of a temporary pond, a narrow temporary stream feeding it and adjacent temporarily flooded grasslands, 0.6 km NNW of the Chingombe catholic mission, near a cemetery; 14°24.90'S, 29°57.80'E, 530 m a.s.l. – the bottom of the rift valley. The pond small (10×12 m), shallow in the beginning of the rainy season and even 1 m deep at the end of it; in 2013 partly and in 2015, totally densely overgrown by bamboos and other grassy and Cyperus-like vegetation.
[13.] Chingombe catholic mission, near a small power station; 14°25.07'S, 29°58.02'E (in the range of 100 m), 520 m a.s.l. – the bottom of the rift valley. Mosaic of open areas with single trees and loose banana plantations, mango woodlands and a dense wet forest patch. With a permanent concrete canal of the power station, an additional temporary shaded canal operating only during high water phases in the rainy season and a small pool in the forest.
[14.] A pool in the fields, 0.4 km SE of the Chingombe catholic mission; 14°25.32'S, 29°58.16'E, 510 m a.s.l. – the bottom of the rift valley. Small, bordered and partly overgrown with dense herbaceous and bushy vegetation.
[15.] The Chingombe River, a road ford 0.95 km ESE of the Chingombe catholic mission; 14°25.22'S, 29°58.51'E, 510 m a.s.l. – the bottom of the rift valley. An open, broadened (to 10–20 m), calm fragment of a small permanent fast clear-water river.
[16.] A pool on the Mulembwe (=Milembwe) temporary stream, 2.3 km NE of the Chingombe catholic mission; 14°24.45'S, 29°59.06'E, 530 m a.s.l. – the bottom of the rift valley. Small and shallow (to >0.5 m). Overgrown with abundant Schoenoplectiella lacustris, but in 2017, after the drought of 2016, only by grassy vegetation. Partly shaded, surrounded by miombo woodlands and grassy areas. As inhabited by very rare Ceriagrion mourae Pinhey, 1969, it was described and illustrated in
[17.] A pool on the Mulembwe (=Milembwe) temporary stream, 0.2. km below the road, 2.4 km NE of the Chingombe catholic mission; 14°24.59'S, 29°59.20'E, 520 m a.s.l. – the bottom of the rift valley. Temporary, small (120 m2), shallow (10–30 cm), with grassy vegetation.
[18.] A pool on the Mulembwe (=Milembwe) temporary stream, 0.4 km below the road, 2.5 km NE of the Chingombe catholic mission; 14°24.65'S, 29°59.30'E, 520 m a.s.l. – the bottom of the rift valley. Temporary, small (75 m2), shallow, with grassy vegetation.
[19.] A pool in the light miombo woodland, 2.7 km NE of the Chingombe catholic mission; 14°24.37'S, 29°59.25'E, 520 m a.s.l. – the bottom of the rift valley. Temporary, small, shallow (to 50 cm), with an open water table bounded with herbaceous (mostly grassy) vegetation.
[20.] The Lukusashi River, 7.9 km SE of the Chingombe catholic mission; 14°27.52'S, 30°01.67'E (both river banks), 435 m a.s.l. – the bottom of the rift valley. Situated in hot open and semi-open landscapes, with bush and woodlands locally on the river banks. Fairly large: 40–200 m wide in the dry and wet season, respectively. Fairly fast flowing with variable water level: shallow and anastomosing within the broad river bed at the end of the dry season and swollen in the rainy season. Bottom deposits are mostly sandy, near banks more gravelly and, in lateral, calm or stagnant branches, sandy-muddy. In the low-water phase, almost without water vegetation, only with rare small patches of Pistia stratiotes and high grass ‘walls’ at the border of water and river banks. In the high-water period, with broadly flooded bank vegetation.
[21.] The Lukusashi River, 8.1 km E of the Chingombe catholic mission; 14°25.50'S, 30°02.51'E, 440 m a.s.l. – the bottom of the rift valley. For the habitat description see [20.].
[22.] Woodlands and a clearing by a ravine, 0.2–0.4 km of the Lukusashi River, 8.1 km E of the Chingombe catholic mission; 14°25.30'S, 30°02.49'E, 450 m a.s.l. – the bottom of the rift valley.
[23.] Two temporary pools (60 m distant) in a large meandering ravine, being a ‘product’ of temporary watercourse erosion, 150–200 m from the Lukusashi River, 8.1 km E of the Chingombe catholic mission; 14°25.38'S, 30°02.50'E, 450 m a.s.l. – the bottom of the rift valley. Situated in semi-open landscape, at the base of the ravine wall. Small, partly shaded by dense bushes with branches overhanging the water table. With soft clayey bottom and turbid shallow water.
[24.] The Bwingi Stream, 5.2 km NE of the Chingombe catholic mission; 14°23.14'S, 29°59.99'E, 510 m a.s.l. – the bottom of the rift valley. Complex of a permanent stream and its tributaries, seepages and temporarily flooded areas in its valley. The main stream mostly shaded with clearings and morphologically strongly diversified between 0.5–1 m wide shallow sections with moderate current and almost stagnant pools to 2 m wide and 1 m deep. In the stream bed, a mosaic of sandy-gravelly bottom and tree roots, abundant detritus and old leaves. The stream headwaters more open and more strongly flowing, with seepages.
[25.] Valley of the Bwingi Stream, 5.2 km NE of the Chingombe catholic mission; 14°23.13'S, 29°59.96'E, 510 m a.s.l. – the bottom of the rift valley. Grassy depressions, partly shaded by loose trees, situated in the fenny areas with seepages feeding the upper reach of the stream. At the very beginning of the rainy season still dry, but may be flooded then as this is suggested by their structure and effects of first intensive rains.
[26.] The Kampoko River, a river ford on the road Chingombe-Mboroma, 26 km NE of the Chingombe catholic mission; 14°14.33'S, 30°07.23'E (to 300 m downstream), 490 m a.s.l. – the bottom of the rift valley. At the very beginning of the rainy season, a small river ranging from 1 m wide watercourse with side puddles up to 10–20 m wide calm sections; mostly shallow, with sandy, gravelly or stony bottom with boulders and some detritus deposits. Partly shaded with trees and bushes overhanging the water, but mostly open.
[27.] A tributary of the Mulungushi River near the Great North Road Kabwe-Kapiri Mposhi, 17.5 km NW of Kabwe; 14°19.10'S, 28°32.81'E, 1,125 m a.s.l. – the plateau. An open small very sluggish river with flooded valley (a few to 20 m wide). With thick soft bottom deposits and water up to 1 m deep, almost completely overgrown with dense herbaceous vegetation.
[16.] 29 January 2017, emergence: 9 teneral individuals (3 M).
[19.] 21 April 2015, emergence: several individuals (2 M, 1 F); 29 January 2017, emergence: several individuals (1 M).
Very widespread in the Afrotropics, from Senegal and Sudan to South Africa, but known from a relatively low number of localities. Also in Zambia, previously known only from few records in the 1960s at the south-eastern border of the country, along the Zambezi River between Livingstone and Katambora Rapids (
[3.] 26–28 January 2017, several individuals, mostly teneral/juvenile, but also adult (2 M: teneral and adult).
[4.] 28 April 2015, 2 adults (1 M).
[5.] 25 April 2015, 1 M, 1 F; 28 April 2015, 1 F.
[6.] 26 January 2017, 4 tandems (2 ovipositing) and 1 female (1 M and 1 F), mostly in the widest (1.5 m) and shallowest, partly sunny section, attracted by favourable oviposition conditions.
[8.] 20 January 2017, 2 males, 1 female (1 M, 1 F).
[11.] 19 January 2017, 1 F.
Widespread in a basin of the mountain Mikwa stream from its source to its lower reach, between 1,180 and 640 m a.s.l. Recorded at six localities, three related to the main stream and three to its tributaries. Occurred at side seepages and trickles and at the main beds of narrow and shallow streams. Teneral and juvenile individuals observed low on the herbaceous vegetation and bushes. Adults settled either quite high (two to several metres) on tree leaves or low in the direct neighbourhood of seepages and trickles. Especially on trees, selected small sunny spots in shady surrounds. Oviposition in submerged tree roots or thin rootlets in a very shallow current water of a small stream (Fig.
Recorded mostly in Zimbabwe, Mozambique and Malawi, with only one locality known in Katanga, DR Congo and one, raising doubts, in western Zambia (Fig.
[1.] 20 December 2013, after first intensive rains but still without water, 3 M and 1 F.
[2.] 30 April 2015, emergence and reproductive activity: 1 tandem, 1 teneral male and 1 adult male (3 M, 1 F).
Mostly West and Central African species reaching south to Zambia and Malawi. In Zambia not rare, but previously known only in northern parts approximately to the latitude of Chingola and the Kasanka National Park (
[1.] 20 December 2013, after first intensive rains but still without water, 2 M.
[12.] 16 December 2013, 1 F.
[25.] In moderate numbers in a very restricted area still without water: 9 December 2013, 2 M; 12 December 2013, 1 F; 17 December 2013, 3 M; 19 December 2013, 3 M, 2 F.
A quite recently described species, previously known only from uplands (1,150 m a.s.l. upwards) in a relatively narrow range between northern and central Zambia and Malawi (
[1.] 20 December 2013, after first intensive rains but still without water, 1 male, 1 tandem (2 M, 1 F); 31 January 2017, several individuals, 1 tandem (5 M, 2 F).
[2.] 30 April 2015, 1 teneral M.
[12.] 18 December 2013, 1 M.
[17.] 20 April 2015, 1 teneral M.
A recently described species, previously known from uplands (1,000 m a.s.l. upwards) in a relatively narrow range from northern Zambia through central and southern Malawi to northern Mozambique (
[20.] 8 December 2013, 3 males (2 M) in two small patches of floating Pistia in lateral river branches (but within the river bed), currently shallow and barely flowing or stagnant, but during the rainy season deeply flooded by the strong swollen river. The species occurrence in such a river is noteworthy as the species is mostly associated with standing and often temporary waters, but also streams (
Mostly occurs in Eastern Africa, from South Sudan to Mozambique (
[12.] 14 December 2013, 3 males above the water table of the pond (2 M).
[14.] 14 April 2015, 1 M.
[15.] 15 April 2015, fairly numerous, tandems (4 M 1 F), a male caught and consumed by male Pseudagrion gamblesi Pinhey, 1978.
[16.] 29 January 2017, 1 M (donated to the collection of Jens Kipping).
[18.] 21 April 2015, 1 teneral M.
[24.] 12 December 2013, 1 teneral M probably originating from the stream.
[26.] 17 December 2013, several males above the water table of wide and calm sections of the river (3 M).
An East African species occurring from Ethiopia to northern South Africa (
[13.] 27 April 2015, 1 M foraging in a small meadow surrounded by trees, during the civil twilight phase.
Mostly recorded between 10° N and 10° S (
[13.] 22 April 2015, 1 M; 26 April 2015, 1 F (Fig.
The female (Fig.
One of most common large Gynacantha-species, mostly known from Eastern Africa and southern parts of Central Africa (
[13.] 15 April 2015, 1 F foraging at various heights, from above the herbaceous vegetation up to several metres, in an open place partly surrounded by trees (in the same place as H. trinervulata in the next days), just before 18:00 h, i.e. shortly after sunset (17:53 h), during the civil twilight phase.
The collected female of Gynacantha sp. (Fig.
i. a quite thin weakly triangular-almost rectangular stem with straight or weakly convex margins (Fig.
ii. a quite flat and rather short upper arm (cross-bar, Fig.
The last trait is not included in keys, but its similarity in the collected male and female of G. villosa and different images in the collected female Gynacantha sp. and male G. vesiculata are worth noticing and checking on larger material.
The COI sequence of the studied female of Gynacantha sp., analysed in BOLD (The Barcode of Life Data System), is:
i. very different from sequences of Gynacantha cylindrata and G. vesiculata;
ii. almost the same as an unpublished sequence of Gynacantha villosa from Tanzania (similarity 99.85%) and very close (99.26%) to a published sequence of villosa from coastal Kenya;
iii. clearly different (similarity 93.83 and 94.29%) from unpublished sequences of Gynacantha villosa from Ethiopia and Zambia.
These results and the BOLD TaxonID Neighbour-Joining Tree suggest that the name Gynacantha villosa comprises two different species and the studied female of Gynacantha sp. from central Zambia belongs to the taxon represented by the Tanzanian-Kenyan sequences. Already
[13.] 11 December 2013, 1 F; 10 April 2015, 1 M, 1 F; 23 April 2015, 1 M; all the individuals foraging in the same very small area of 400 m2; on 10 April (sunset 17:57 h, end of civil twilight 18:18 h) the female appeared at 17:46 h flying very low in a deep shade under the mango tree while the male was collected at 18:00 h in a slightly lighter place but still under trees.
As only few individuals of the species were earlier known and analysed (
A new species for Zambia. Previously known only from four localities, two in E Tanzania (Kichi Hills and Muheza District), one in N Malawi (Nkhata Bay) and one in Katanga (Lubumbashi) in DR Congo (
[3.] 26 January 2017, 2 ovipositing females (1 F, Fig.
Two females: very large, Hw 54.2 and 55.3, FwAx 31 and 31/30, FwMx 6/7 and 5(and two halves)/7, HwMx 5 and 6. In one female FwMx 1–2 higher than HwMx in both wings, in the second female such a situation in one wing. HwAl 12/14 and 12/11. Measurable and countable data on female fuliginosa are scarce. However,
A new species for Zambia. Mostly recorded from Western and Central Africa, with only few records south of 5° S and two localities known from Katanga in DR Congo (
[13.] 16 April 2015, 1 F; 18 April 2015 1 M; 23 April 2015 1 F; all the individuals juvenile, their condition suggested their emergence on the same day. Individuals observed just before a thunderstorm on 17 April 2015 and well before sunset on 27 April 2015 highly resembled those collected (size, colours, behaviour) probably also being trinervulata. All the individuals foraged at moderate to fairly great heights, from two to several metres, in open and semi-open places in a small area of several hundred square metres and were present 13–18 minutes before sunset (time of record/sunset): 17:40/17:53 h, 17:35/17:52 h, 17:31/17:49 h.
Due to juvenile condition, colouration of individuals is partly useless. However, all the substantial diagnostic traits, i.e.: a) size (Hw 35–37), b) details of relatively open venation (Fig.
The Zambian individuals were distinguished by lower numbers of cross-veins versus higher numbers given by
Due to poor and not always clear material,
Previously known only from upper Zambezi rapids in north-westernmost corner of Zambia (
Heliaeschna trinervulata from Chingombe, Zambia: male (a–d) and female (e), with recognisable diagnostic morphological traits such as: relatively open venation (a, e) with a low number of cross-veins (e.g. Ax and Mx), the pale frons with a barely marked cross-bar of T-mark (b, e), wing bases without dark subcostal rays (a, e) but with distinct amber tint in the female (e) and shape of male cerci (c, d) (phot. B. Daraż).
[21.] 15 December 2013, 1 exuviae near the bank in the transitional zone between the main flowing river branch and the side calm arm fed by seepages. Its diagnostic traits, i.e. the total length 36.5, the convex anterior border of prementum, the finger like crenation of the labial palpus, proportions between the leg length and abdomen size, as well as between the tube (S10) length and the total length, were typical of the genus and almost exactly the same as illustrated by
Historically, true M. dissimilis was known only from an area between 15° and 20° S, from three localities in S Malawi, E Zimbabwe and at the southern border of Zambia (locus typicus Katambora rapids of the Zambezi River) (
[9.] 7 December 2013, 1 teneral F (Hw 31, Abd 35.5, pt 3.5; Fig.
The colour pattern already sufficiently developed to clearly assess the size and shape of pale and dark areas. The following features of this female (partly visible in Fig.
Several other features of the studied female may, however, raise some doubts about species identification. They are given below versus their different picture in
The middorsal area of the thorax appears the most problematic. In the studied female, the whole middorsal area along both the middorsal carina and a proximal section of the ante-alar ridge is pale; only the dorsal edge of the proximal ante-alar ridge is darkened (Fig.
The COI sequence of the studied female, analysed in BOLD (The Barcode of Life Data System), appears quite close to known unpublished sequences of Notogomphus zernyi, with similarity 98.29%. The mean genetic distance between COI sequence groups recovered in the Neighbour-Joining analysis is 1.85% (SD=0.51) (the analysis due to K.D.-B. Dijkstra pers. comm.).
Considering the middorsal problematic feature, the not-completely matched species level by the molecular data and the tendency in this genus towards forming separate taxa in isolated areas, a new species of Notogomphus cannot be totally excluded. However, the vast majority of features, typical of the species or covered by the intraspecies variation, as well as the quite high similarity of COI sequences, suggest the female of Notogomphus zernyi. Therefore, we have decided to qualify it as cf. zernyi.
N. zernyi is known from E Zimbabwe, Malawi and SW Tanzania (
Notogomphus cf. zernyi, a teneral female from the Mikwa Stream, near Chingombe, Zambia, with details of the thoracic pattern in dorsal and lateral views (a–c), a subgenital plate (d) and the pattern of the proximal segments of abdomen in dorsal view (e); arrows indicate the pale distal part of the middorsal stripe (phot. B. Daraż).
[3.] 26 January 2017, 3 males (2 M); 27 January 2017, 1 F; 28 January 2017, 1 M. Apart from a male intensively flying back and forth on the adjacent semi-open slope, all other individuals were observed above the water, mostly in shaded places where males flew rapidly.
In the collected males, an epiproct is only slightly shorter than cerci and all the males and the female are almost uniform in their dark and yellow pattern. A yellow spot in the frons groove and prejugal spots on S3–5 (on S5 small), separated by a black median line, point to an intermediate form (close to former onerata) between dark and pale extrema of the species (see
Occurs in Eastern Africa, southwards reaching the northern parts of South Africa; its range also includes southern parts of Central Africa, i.e. Katanga and Zambia. In Zambia, previously known from several old records in the north-western ‘corner’ of the country around Ikelenge-Mwinilunga (
[3.] 27 January 2017, 3 territorial males and 1 tandem in copula (Fig.
[7.] 26 January 2017, 2 territorial males (1 M collected and immediately replaced with the second one).
[10.] 19 January 2017, 2 males: an aggressive resident and a visiting intruder (with a clash); 20 January 2017, 1 territorial M.
Relatively widespread in a basin of the Mikwa mountain stream, from its source to lower reach (1,180–640 m a.s.l.). Related to very small pools in the stream valleys and wider calm sections of streams in their headwaters. All the four places were: small (a few m2), relatively deep (0.3–1 m), generally calm, but with some invisible flow or at least with water supply, permanent or at least semi-permanent, shaded with clearings, with tree branches overhanging at a height of several metres, with a layer of detritus and fallen leaves on the bottom, with sticks or stones emergent from the water or protruding from the shore and serving as roosting sites low above the water table. The male behaviour very consistent. Smaller pools ‘housed’ only one territorial male chasing away appearing intruders while, in the largest pool (a widened section of the stream), two residents occurred at a safe distance of 2.2 m. Although roosting sites were most frequently shaded, all the males appeared only in sunny weather and disappeared in cloudy times. Very shy: each movement in the surroundings was followed by their immediate, rapid and surprisingly unobtrusive vertical escape flight into the tree branches zone (see also
A typical East African species, mostly occurring east of 30° E and ranging from Uganda and Kenya to South Africa (
[23.] 21 January 2017, 3 territorial males (2 M) on overhanging dry bush branches and on sticks 0.5–2.5 m above the water table of pools. One of them also with a female in copula; the copulation wheel was formed in flight and the pair in this position flew down between overhanging bush branches. Temporary nature of the pools, their location in a ravine and presence of overhanging branches and soft bottom highly resemble the habitat description given by
A new species for Zambia although two nearest known localities are situated in Zimbabwe just at the border with Zambia (
[1.] 31 January 2017, 1 male patrolling large areas above the water table at a distance from the shores and 1 tandem: not collected but undoubtedly identified.
[2.] 30 April 2015, 1 male persistently patrolling above the water table, at a distance from the shores (1 M).
The species with a giant Afro-Asian distribution range, widespread in the Afrotropics and locally common, e.g. in parts of South Africa, but surprisingly rarely recorded in Central Africa (
[26.] 17 December 2013, 3 M in sunny places with stones and boulders at broader, both calm and more flowing sections of the river.
An East and South African species (
[27.] 13 January 2017, several males and 1 tandem (2 M).
Occurs mainly in Central Africa (
[22.] 21 January 2017, several males (3 M, two immature); 24 January 2017, in moderate numbers, dispersed at several sites. Perched in sunny places on bushes and especially on tips of sticks above the herbaceous vegetation in woodlands and at clearing edges. Certainly associated with the Lukusashi River (the locality [21.]), but absent from the bank vegetation and direct neighbourhood of the swollen river. Such a situation, noticed as often occurring in this species (
A largely East African species penetrating here and there into central and south-western parts of the continent (
Our studies revealed that distribution ranges of several species, such as Aciagrion africanum, Gynacantha vesiculata, G. immaculifrons, Heliaeschna fuliginosa, H. trinervulata and Trithemis nuptialis, reach further south that was previously known (Fig.
Distribution of selected dragonfly species in the southern Afrotropics (between the Equator and the southern border of Zimbabwe): a Heliaeschna fuliginosa b Gynacantha vesiculata c Heliaeschna trinervulata d Gynacantha immaculifrons e Notiothemis jonesi f Allocnemis marshalli g Notogomphus zernyi h Mastigogomphus dissimilis. The black dot – the new locality. Other points – other data according to literature and Odonata Database of Africa (J. Kipping pers. comm.,
The evening concentration of five species of Gynacanthini in a small area near the power station in the Chingombe mission showed especially favourable conditions existed there, at least for foraging activity. Although larval habitats were not found, the flight of Gynacantha immaculifrons in subsequent years exactly under the same tree, as well as the presence of juvenile Heliaeschna trinervulata, suggested the native occurrence somewhere nearby, possibly in an adjacent patch of very dense wet forest and bush. This concentration occurred at the end of the rainy season, but, in the peak of it (in January), imagines of Gynacanthini were not recorded there despite special searches. Both of these facts, as well as colours and state of collected imagines, suggested that this concentration was related to the immature phase of life.
Crepuscular activity was typical of all the five species of Gynacanthini recorded in the mission. However, the measured data and several unmeasured but localisable-in-time situations showed a sequence of species. Heliaeschna trinervulata appeared as the first, still in quite good light conditions, beginning from the latest sunshine. In contrast to Pinhey’s observations from Malawi (1966), it was not recorded at dusk. It was flying in more open, unshaded conditions at moderate to fairly great heights similar to those given by
Although precise data on habitat selection by gynacathines are generally scarce, species of Heliaeschna are perhaps associated with more permanent water than Gynacantha, like swamp forest or streams (
We are grateful to Jens Kipping and Klaas-Douwe B. Dijkstra for valuable remarks and information, to Bertie Brink and Herb Kageler for their recent data on Notiothemis jonesi and Gynacantha immaculifrons and to Miroslawa Dabert for help in the obtaining and analysis of molecular data. These studies would not have been possible without the warm hospitality and far-reaching help of the late unusually good man, Reverend Marceli Prawica from the Chingombe Mission.