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Africa's smallest damselfly - A new Agriocnemis from Namibia (Odonata: Coenagrionidae)


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Agriocnemis bumhilli sp. n., a new damselfly from the Kwando River in northeastern Namibia is described. The new species is similar to Agriocnemis angolensis but characterized by unique male appendages, swollen abdominal segments 9 and 10, the complete absence of antehumeral stripes, and smaller size. The species is illustrated and a photograph is provided. For comparison, an illustrated key to the other members of Agriocnemis within south-central Africa is provided.
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Africas smallest damselflya new Agriocnemis
from Namibia (Odonata: Coenagrionidae)
Jens Kipping & Andreas Martens & Frank Suhling
Received: 20 October 2011 / Accepted: 14 March 2012
Gesellschaft für Biologische Systematik 2012
Abstract Agriocnemis bumhilli sp. n., a new damselfly
from the Kwando River in northeastern Namibia is de-
scribed. The new species is similar to Agriocnemis ango-
lensis but characterized by unique male appendages,
swollen abdominal segments 9 and 10, the complete ab-
sence of antehumeral stripes, and smaller size. The species
is illustrated and a photograph is provided. For comparison,
an ill us tr ated key to the othe r members of Agriocnemis
within south-central Africa is provided.
Keywords Damselfly
Agriocnemis taxonomy
Body size
Kwando River
Measurements and morphology
Fw Forewing(s)
Hw Hindwing(s)
Pt Pterostigma
Ax Antenodal cross-vein(s)
Px Postnodal cross-vein(s)
S110 Firsttenth abdominal segment
Natural History Museums
NMNW National Museum of Namibia Windhoek
RMNH Netherlands Centre for Biodiversity Naturalis
The term cerci is used for male superior anal appendag es the
term paraproct for the male inferior anal appendages. The
gender symbols ( for male, for female) are used when
The genus Agriocnemis Selys, 1877 consists of 41 species,
which occur in Africa and Asia (Pinhey 1974, Schorr &
Paulson 2012). The members of the genus are characterized
by the position of the arculus far distal to the second Ax, and
their small size (Pinhe y 1974). Some species belong to the
smallest known zygopterans. In his revision, Pinhey (1974)
listed 16 species of Agriocnemis for the African continent
and adjacent islands. Agriocnemis aligulae Pinhey, 1974
was later synonymised with A. maclachlani Selys, 1877
(dAndrea & Carf i 1997, vide Dijkstra 2007a). Some of
the various subspecies described by Pinhey (1974 ) remain
dubious and one, Agriocnemis pygmaea sania, was raised
back to species level, A. sania Nielsen, 1959 (Dumont
1974). In recent years, dAndrea & Carfi (1997) described
A. dissimilis from a single female, which, however,
appeared to be a synonym of the Ugandan endemic A.
palaeforma Pinhey, 1959 (Dijkstra 2007a).
Of the seven species of A
iocnemis known for the southern
Africanregion(Suhlingetal.2009), six occur in the Zambezi
and Okavango River basins, namely A. angolensis Longfield,
1947, A. exilis Selys, 1872, A. gratiosa Gerstäcker, 1891, A.
pinheyi Balinsky, 1963, A. ruberrima albifrons Balinsky, 1961
This paper is dedicated to Prof. Dr. Michael L. May in honour of his
work on Odonata taxonomy.
This is a contribution to the Festschrift for Michael L. May.
J. Kipping (*)
Naturkundemuseum Mauritianum Altenburg,
Parkstrasse 1,
04600 Altenburg, Germany
A. Martens
Pädagogische Hochschule Karlsruhe,
Bismarckstrasse 10,
76133 Karlsruhe, Germany
F. Suhling
Institut für Geoökologie, Technische Universität Braunschweig,
Langer Kamp 19C,
38106 Braunschweig, Germany
Org Divers Evol
DOI 10.1007/s13127-012-0084-4
and A. victoria Fraser, 1928 (Mart ens et al. 2003,Kipping
2010, Pinhey 1984). Most of the species are to be
found i n the extensive swampy wetlands of permanent
rivers such as the Okavango Delta and Linyanti
swamps. The South African endemic A. falcifera
Pinhey, 1959 is restricted to m ountainous habitats.
In February 2006, A.M. collected a number of
Agriocnemis species at the type locality Kwando River
at Bum Hill (see Fig. 4). With many A. exilis and A.
victoria, two males were collected that were noticed as
unusual. In Suhling & Martens (2007:p.92)thetwo
specimens were illustrated erroneously on a photograph
under A. angolensis. At this stage, it was already clear
that the specimens had some characters unusual in
Agriocnemis. Therefore, during a fieldtrip in January
2009, J.K. also visited the type locality with the aim of
collecting more specimens. Here, we describe the new
species formally based on seven males and four females.
Agriocnemis bumhilli sp. n.
(Figs. 1, 2)
Suhling & Martens (2007: photograph p. 92)
Agriocnemis angolensis nec Longfield, 1947.
Etymology The species is named after its type locality, the
community campsite Bum Hill on the banks of the Kwando
River, Namibia. The name also nicely refers to one of the most
prominent characters of the species, the swollen S910.
Material examined Holotype : Namibia, Caprivi Region,
Kwando River at Bum Hill community campsite, northwest
of Kongola (17°4653S, 23°2027E, 969 ma.s.l.), 10
January 2009, leg. J. Kipping, to be deposited in
NMNW.Allotype and paratypes (all from the same local-
ity): 6 ,4, same locality and date as holotype, allotype
female in copula with paratype male (2 preserv e d in
100% ethylalcohol, a ll others acetone dried in plastic enve-
lopes); paratype and 2 paratype to be deposited in
NMNW, 2 paratypes in RMNH and others in coll. J.
Kipping.Other material: 2 , 28 February 2006 leg. A.
Martens, to be deposited in NMNW, 1 , 7 December 2010,
leg. F. Suhling, to be deposited in NMNW.
Description of holotype male
Acetone dried
, preserved dry, in plastic envelope.
Head Labium whitish. Mandibles pale green. Labrum black
with broadly pale ventral margin. Postclypeus black, the
anteclypeus contra sting pale green. Genae and antefrons
pale green. Frons, vertex and occiput black with bronze to
green metallic sheen. Frontal band severed. Base of anten-
nae and ocelli pale brown. Isol ated postocular spots small,
slightly elongated and bright blue. Postgenae pale green
with contrasting black markings. Dorsal half of eyes in life
black, sharply separated from pale green ventral half.
Thorax Prothorax dorsally black with green met allic sheen.
Anterior collar black without markings. Lower part of the
sides pale green. The prothoracic hindlobe bears only a
broad and flat middle lobe with a thin pale line along
Fig. 1 ad Agriocnemis bumhilli sp. n.holotype male. a Prothorax
and thorax, lateral view. b Distal portion of abdomen, dorsal view. c
Appendages, posterior view. d Distal portion of abdomen, lateral view.
Drawings by Ole Müller
Fig. 2 Agriocnemis bumhilli sp. n.male, resting in dense vegetation
at t he Kwando River, Bum Hill, Cap rivi Strip, 10 January 2009.
Photograph by J. Kipping
J. Kipping et al.
hindmargin. Lateral lobes not developed. Laminae black
with pale green at distal end. Synthorax completely black
with strong metallic sheen down to interpleural suture. No
pale antehumeral stripes. Small black spot on metapleural
suture. Sides pale green.
Legs Coxae and trochanters pale. Femora pale with a sharp
brown stripe on exterior surface that broadens distally and
forms a brown ring at the distal end. Tibiae all pale with a
thin exterior dark stripe and contrasting dark setae. Tarsi
pale with dark ring at distal ends of segments.
Wings Clear, venation dark brown. Fw with 7 Px, in Hw 6
Px. Pt in all wings pale framed by thick dark brown margins.
Abdomen with continuous broad black do rsal band a nd
bronze sheen on S17. S13 pale green at sides changing
to bluish on S4.7. S8 10 with purplish color ation. S9 and
S10 remarkably swo llen both in lateral and dorsal view but
distal end of S10 not raised apically. Anal appendages
purple orange. Superior appendages longer than S 10. In
lateral v iew broad, parallel sided in the distal half and
pointed apically. In dorsal view broad at base and also
becoming point ed apically. At the base of each of the supe-
rior appendages a downward directed flat and pointed pro-
cess, the a pical tooth of which reaches the inferior. A
remarkable tuft of hairs on this process resembles a back-
ward directed pointed structure. Inferior appendages small,
blunt and with a small dark upward-directed tooth.
Measurements [in mm] Total length 18.4 (incl. anal appen-
dages), abdomen length 14.6 (excl. anal appendages), head
width 2.5, Fw length 9.4, Hw length 8.8, Pt in Fw 0.6.
Description of female
Collected in copula with a paratype male. Acetone dried,
preserved dry, in plastic envelope.
Head Labium whitish. Mandibles bluish white. Labrum
black with broadly pale ventral margin. Postclypeus black,
the anteclypeus contrasting pale green. Genae pale green,
antefrons mainly brownish black. Frons, vertex and occiput
black with dull green metallic sheen. Base of antennae and
ocelli pale brown. No postocular spots. Postgenae whitish
without markings. Dorsal half of eyes in life black, sharply
separated from pale green ventral half. Traces of pruinosity
at sides of anteclypeus.
Thorax Prothorax dorsally black with green met allic sheen.
Anterior collar black without markings. Lower part of the
sides pale blue. The prothoracic hindlobe narrow, slightly
erect and not remarkably shaped. Lateral lobes not devel-
oped. Laminae black, pale blue at distal end. Synthorax
completely black with strong metallic sheen down almost
to interpleural suture. No pale antehumeral stripes. Small
diffuse brown spot on second lateral suture. Sides pale
bluish green. Slightly pruinose on prothorax.
Legs Coxae and trochanters pale. Femora pale with a sharp
brown stripe on exterior surface that broadens distally and
form a brown ring at the distal end. Tibiae all pale with a
thin exterior dark stripe and contrasting dark setae. Tarsi
pale with dark ring at distal ends of segments.
Wings Clear, venation dark brown. Fw with 8 Px, in Hw 6
Abdomen with continuous broad black do rsal band and
bronze brown sheen on all segments and pale blue at sides.
S710 with a brown hue.
Measurements [in mm] Total length 18.5 (incl. anal appen-
dages), abdomen length 14.2 (excl. anal appendages), head
width 2.5, Fw length 11.1, Hw length 9.9, Pt in Fw 0.6.
A dark Agriocnemis similar to A. angolensis and A. exilis in
general coloration but with a darker thorax without a trace of
antehumeral stripes. It is easily distinguished from all other
species of the genus by the unique shape of the male
appendages and the male swollen abdominal S9 and S10
(Fig. 1). Other Agriocnemis species never show such swol-
len last abdominal segments (Fig. 3). The complete absence
of antehumeral stripes is also unusual in the genus.
Body size Total length ranges from 17.7 to 19.2 mm (incl.
anal appendages). Length of abdomen range from 13.6 to
14.6 mm (excl. anal appendages). Length o f Hw range from
8.5 to 9.1 mm.
Prothorax T
males show pale bluish markings on the
anterior collar in form of a small dot at each side of the
collar, in one male there is only a tiny dot on the left side.
All the other males have complete black anterior collars.
Postgenae The rear of the head is white in all males. Some
show a distinctive and sharp dark brown to black pattern,
others just a diffuse brown hue.
Africas smallest damselfly
Frontal band The pale frontal band is always severed by a
dark central line that links the dark coloration of the post-
clypeus with that of the frons. The width of this line varies
from a narrow line to a broad band that is half as wide as the
Wings One male has a malformation with a remarkable
short left Fw with only 8.0 mm in length and only 1 Px
Body size Total length range from 18.5 to 19.5 mm (incl.
anal appendages). Length of abdomen range from 14.2 to
16.2 mm (excl. anal appendages). Length o f Hw range from
9.9 to 10.4 mm.
Head Only one specimen shows small and pale blue post-
ocular spots.
Diagnosis of Agriocnemis species of the region
A key to the Agriocnemis males of the Zambezi and
Okavango River basins is presented. Please note, this key
is included only for the designated region, outside this area
it could lead to mi sidentification. Other similar species
occur in South Africa and in the tropical forests and swamps
north of the Zambezi basin.
1 Cerci much longer than paraprocts; cerci as long or
longer than S10 (i.e., Fig. 3a-d) ................................. 2
- Cerci of equal length or shorter than paraprocts; cerci
shorter than S10 (i.e. Fig. 3gi) ................................... 4
2 Cerci narrow and forcipate, in dorsal view curved to-
wards each other (Fig. 3a,b); thic k black stripe along
metapleural suture ....................................... A. victoria
- Cerci broad, not forcipate; black stripe along metapleu-
ral suture reduced to a black dot at dorsal end .......... 3
3 Antehumeral stripes absent; abdominal segments S9
and S10 swollen; cerci broad at base and tapering to a
Fig. 3 ak Morphological
characters of the other members
of Agriocnemis in south-central
Africamale abdominal
appendages. a,b A. victoria lat-
eral (a) and dorsal (b) view. c,d
A. angolensis lateral (c) and
dorsal (d) view. e,f A. gratiosa
lateral (e) and dorsal (f) view. g
A. ruberrima lateral view. h A.
pinheyi lateral view. i A. exilis
lateral view. j,k male prono-
thum, dorsal view: j A. pinheyi;
k A. exilis. Drawings by
K.-D.B. Dijkstra
J. Kipping et al.
pointed apex (Fig. 1ad). Pt of Hw not with broadened
costa ............................................................ A. bumhilli
- Antehumeral stripes normally developed; last abdomi-
nal segments not swollen; cerci spatulate, in dorsal view
parallel sided with rounded end (Fig. 3c, d); Pt of Hw
with distinctly broadened costa ............... A. angolensis
4 Cerci short er than paraprocts, slender paraprocts taper-
ing to single points, at most 2x as long as cerci (Fig. 3e,
f). Hw length more than 11 mm ................... A. gratiosa
- Cerci about as long as paraprocts; smaller species, Hw
length usually less than 10 mm ....................................5
5 ApexS10raiseddistinctlytoanuprightprocess
(Fig. 3g)...... ........................................... A. ruberrim a
[ssp. A. r. albifrons (N-Botswana, NE-Namibia) with
frons whitish pruinose, S17 bro adly black, S810
orange-red; ssp. A. r. ruberrima (NE-South Africa,
coastal Mozambique) abdomen mainly orange-red,
frons not pruin ose]
- Apex S10 hardly raised .............................................. 6
6 Hindlobe of prothorax incised at two sides, separating a
fan-like middle section (Fig. 3j); cerci without ventral
needle-like process (Fig. 3h) ......................... A. pinheyi
- Hindlobe of prothorax not incised (Fig. 3k); cerci with
ventral needle-like process (Fig. 3i) ................. A. exilis
Habitat and ecology
Agriocnemis bumhilli is known only from the type locality,
although several surveys were carried also out in other parts
of the Kwando River in N amibia and Botswana. Thus,
knowledge of ecological preferences is limited. At Bum
Hill, all the individuals were found exclusively at the margin
of the main channel of the river where a narrow band of
dense grassy vegetation was growing in and along the
running water. A. bumhilli was fou nd there togethe r
with a few indivi duals of A. gratiosa an d A. victoria.
At the adjacent backswamp, A. victoria was the most
abundant Zygopteran, and also A. exilis and A. ruber-
rima albifrons werefoundinlowernumbersinJanuary
and February. In December, only one male of A. bum-
hilli was recorded, while A. ruberrima albifrons was the
most abundant Agriocnemis in the area. According to
the few records the flight season of A. bumhilli is from
the beginning of December to March.
Fig. 4 Map of southern Africa showing the type locality of A. bumhilli sp. n. (white circle) and the density of species of the genus according to the
inferred distribution maps created by the IUCN Red List Assessments (Dijkstra et al. 2011; Suhling et al. 2009)
Africas smallest damselfly
The new species is another example of the great diversity of
anal appendages in t he genus noted by Pinhey (1974).
Agriocnemis bumhilli would fit into group 2 of the prelim-
inary grouping of African Agriocnemis species by Pinhey
(1974), and would stand together only with A. angolensis,
which is also characterized by long and narrow cerci. But it
differs notably from this species in some characters. A.
angolensis has a very remarkable Pt in the hindwing with
a widened and pruinosed marginal edge, a feature unique in
all African Odonata. In the male, the abominal S9 and S10
of A. angolensis are not swollen and both sexes also have
well developed antehumeral stripes. So, if following the
proposed grouping of Pinhey (1974) the new speci es would
consequently need its own group due to these very signifi-
cant distinguishing features. This example shows t hat a
grouping within the genus is pointless.
The body dimensions of the smallest collected male with a
total length of only 17.7 mm, an abdomina l length of 13.6 mm
and Hw length of 8.5 mm makes A. bumhilli to the smallest
member of the genus and of all African Odonata. In comparison
to other species (vide Pinhey 1974) and to our own measure-
ments of specimens collected within the region and at the type
locality at least, this one male of thetypeseriesisprobablyone
of the smallest Odonata specimen ever measured in Africa.
Similar in small body size in worldwide asp ect are some speci-
mens of the Asian Agriocnemis nana (Laidlaw, 1914) with
abdominal length of 14.2 mm and Hw length of 7.5 mm (coll.
R. Garrison) and of Agriocnemis minima Se lys, 1877 from Java
with abdominal length 13.5 mm and Hw length of 8.5 mm.
So far, the Kwando River and swamps around Bum Hill
with five species is the place with the highest diversity of
Agriocnemis species in Africa known from a single locality
overall. Going to the wider scale of the sub-basin catch-
ments with A. angolensis, a sixth species occurs in the area.
The map in Fig. 4 shows the species density of Agriocnemis
in southern Africa. The map is based on inferred distribution
maps created by the IUCN Red List Assessments (Dijkstra
et al. 2011; Suhling et al. 2009). They combine point local-
ity data and expert knowledge on the level of river sub-
basins delineated by the HYDRO1k Elevation Derivative
Database for Africa (USGS EROS).
All the species of the region belong to the group of open
swamp species, most of them distributed widely on the
African continent. But with Agriocnemis angolensis, A. bum-
hilli and A. ruberrima, the south-central Zambezian Odonate
fauna hold some species restricted to the biome. As in the
genera Aciagrion, Trithemis and Pseudagrion, the Zambezian
region with its geological complexity and climatic history
form a recent speciation center also for swamp-dwelling
Agriocnemis (vide Dijkstra 2007b; Dijkstra et al. 2011).
When recognising the Pan-African scale, only the border
on of eastern DRC and Uganda have more Agriocnemis
species in some sub-basins.
Acknowledgments The authors are very grateful to Ole Müller and
K.-D.B. Dijkstra for producing the drawings. H. Fliedner gave some
advice on latin grammar for scientific names.
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... In both regions, recent explorations have led to the discovery of new species (Dijkstra et al., 2007;Ferreira et al., 2014), new populations of threatened endemics (Chelli et al., 2019) and other regionally rare species (e.g. Kipping et al., 2012;Deacon and Samways, 2017;Khelifa and Zebsa, 2018). These important findings suggest that there is great merit in additional exploration to find new odonate species, even in areas with comparatively high sampling effort . ...
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This study has been conducted aim to determine the harmful and beneficial insect species in paddy cultivation in Çanakkale and also identifying the most common weed species of paddy crop in Çanakkale region as well as in the world. The research has been carried out in paddy growing villages of Biga (Savaştepe and Kuruoba villages) and Ezine (Akköy and Güllüce villages) Districts of Çanakkale Province during the paddy cultivating season in 2019. Sampling has been done twice in a month between May and October. Collection of insect samplings was made by taking into account the vegetative development periods of the paddy crop. Accordingly, a total of 40 samples have been taken, at least twice, during the vegetative stage (0-30 days), generative stage (30-60 days) and delivery stage (60 days and onward) of the paddy crop. In other words, a total of 10 samplings have been taken from each location. Insect species have been collected by using netting/sweeping method and then the collected insect samples were brought to the Biological Control Laboratory of the Faculty of Agriculture, Department of Field Crops for further taxonomic studies. The collected insect species have been classified whether they are beneficial, harmful and beneficial/harmful to the paddy crop, and then their morphological, physiological and biological information have been revealed. According to the results of this study; a total of 20 different insect species were identified in the villages of both districts. It has been determined that 8 of these insect species are beneficial, 7 of them are harmful and 5 of them are beneficial/harmful to the paddy crop. Generally, the mode of action of these insects either dirctly cause damage or transmit diseases to the root, stem, leaf, spikes and/or seed parts of the paddy crop. Generally, in order to minimize the infestation of these insect species in the paddy crop, different control methods such as cultural, mechanical, biological, biotechnological, chemical and integrated pest mangement programmes are being applied worldwide. As a result of this study, it is recommended to apply the above mentioned control methods against the infestation of insect pests and weeds that cause major financial losses in paddy cultivated areas. It is also recommended to minimize the pesticides application for the protection and sustainability of beneficial insect population in paddy growing areas.
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In this paper the trends of dragonfly expansions during the last decades in Germany and Europe are summarized. It is shown, that there is a general expansion of many species to the north: Mediterranean species expanded to Central and Northern Europe, whereas some African species expanded to Southern Europe, some are even new to the continent. In general this means an increase of biodiversity, but looking at the ecological effects, in the medium term a decrease can be expected for mooreland and alpine species. Dragonflies can be regarded as a good indicator group for climatic change. Already now in some areas or regions negative effects on waters bodies and their dragonfly communities can be observed and more will occur if e.g. temperature rises or precipitation decreases. The consequences for nature conservation strategies – such as the NATURA 2000 network – are outlined and the general need for monitoring programmes is emphasised.
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In nature, both similar and disparate sizes of Libellula lydia and L. luctuosa larvae frequently co-occur in time and space. To determine if these larvae interact as competitors, and/or predators and prey, I used artificial ponds to manipulate density, species composition, and size range of co-occurring larvae. Detailed life history data were used to design separate fall and spring experiments. In both experiments, @'competition treatments@' contained only larvae similar in size, whereas @'predation treatments@' contained larvae disparate in size. In fall competition treatments there were no density-dependent growth responses. However, in the spring experiment, larvae of both species grew significantly faster in low density than in high density treatments. This seasonal difference in competition was attributed to fluctuations in resource abundance. Competition did not directly affect survivorship. In spring and fall predation treatments, mortality was significantly higher than it was in competition treatments. Inter-odonate predation accounted for 25-45% of total larval mortality in fall, but only 10-15% in spring. In the absence of inter-odonate predation, total mortality was lower for larger larvae than smaller larvae, suggesting the latter are more susceptible to predation by other invertebrate predators. Thus, competition, by decreasing growth rates, should indirectly affect larval survivorship. These data provide evidence that competition and predation will simultaneously affect coexistence between these two dragonfly species. Predation early in larval development should ameliorate the intensity of subsequent competitive interactions at a time when resources are most likely to be limiting. This type of mixed competition/predation interaction is analogous to predator-mediated coexistence, and might explain how such ecologically similar species can coexist at such high densities.
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Kurzfassung Kipping, J.: Die Libellen Botswanas – eine kommentierte Checkliste mit Angaben zu Verbreitung, Phänologie, Habitaten und Rote Liste Status der Arten (Insecta: Odonata). Zwischen den Jahren 2000 und 2010 wurden in Botswana und dabei besonders im nördlichen Landesteil 111 Libellenarten nachgewiesen. Gemeinsam mit weiteren publizierten Funden und Nachweisen ande-rer Sammler vergrößert sich damit die Checkliste der Libellen Botswanas auf 127 Arten. Davon sind 21 Arten neu für die Fauna des Landes. Die hier vorgelegte Checkliste liefert für jede Art Angaben zur Verbreitung, der Phänologie, den besiedelten Habitaten und dem aktuellen Rote Liste Status des IUCN. Die einzelnen Nachweise werden detailliert aufgeführt, im Anhang werden die vom Autor selbst untersuchten Fundorte beschrieben. Die Verbreitungsmuster der Libellenarten in den einzelnen Süßwasser-Ökoregionen werden diskutiert. Abstract Between 2000 and 2010, 111 species of Odonata have been recorded mainly from northern Botswana. Together with other published and unpublished records, this increases the checklist of the Odonata of Botswana to 127 species, of which 21 species have not been recorded before in Botswana. This updated checklist provides information for each recorded species on distribution, phenology, preferred habitats and specific notes, where appropriate. A detailed list of records is given, the localities sampled by the author are described in the appendix. The Red List status according to the IUCN assessment is given. Distribution patterns of the Odonata species in different freshwater ecoregions are discussed.
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The trends of 60 Dutch dragonfly species were calculated for three different periods (1980–1993, 1994–1998 and 1999–2003). Comparing period 1 and period 3 shows that 39 of these species have increased, 16 have remained stable and 5 have decreased. These results show a revival of the Dutch dragonfly fauna, after decades of ongoing decline. The species were categorized in different species groups: species with a southern distribution range, species with a northern distribution range, species of running waters, species of fenlands and species of mesotrophic lakes and bogs. The trends of these different species groups were compared with the all-species control group. As expected, a significantly higher proportion of the southern species show a positive trend than the all-species group. In the northern species group on the contrary, a significantly higher proportion of the species show a negative trend than the all-species group. Different explanations for these results are discussed, such as climate change, improved quality of certain habitats and degradation of other habitats. It is likely that the observed increase of southern species is at least partly caused by the increasing temperatures. The less positive picture of the northern species group is probably more influenced by other environmental factor than directly by climate change. Three out of six southern species which have become established since 1990 have done so during the aftermath of large invasions. It is concluded that dragonflies are well capable of using changing climate circumstances to colonise new habitats.
Since the beginning of this century there have been substantial declines in the distribution and abundance of native Megalagrion damselflies on the Hawaiian Island of Oahu. Native damselflies have also vanished from most low elevation areas on other Hawaiian Islands, although historically, lotic and wetland dwelling damselfly species were once common throughout the archipelago. It is hypothesized that poeciliid fish introduced for biological control have caused the decline of four stream-breeding damselfly species on Oahu, and the extinction or near-extinction of two other species in Hawaii. This study documents the presence of remnant Megalagrion populations in Oahu streams, wetlands and estuaries, and records the elevational distributions of introduced fish in each waterbody surveyed. The distributions of introduced Odonata are also recorded, because the seven species of damselflies and dragonflies introduced to Oahu since 1936 present another potential threat to native Hawaiian damselflies. Native damselfly and introduced poeciliid fish distributions were mutually exclusive on Oahu, and it is concluded that this is probably due to predation by the introduced fish. By contrast, even the rarest native Megalagrion damselflies were found in areas containing introduced damselflies and dragonflies.
In Israel and Sinai Agriocnemis sania Nielsen occurs and not Agriocnemis pygmaea (Rambur). A. sania was previously known only from the oasis of Gat (Libya), and was believed to be endemic to that area. Its original description is revised and taxonomic, ecological and biogeographical data are presented.
Ethograms of intraspecific agonistic behaviours are presented for middle (6–8th) and late (12–14th) instar larvae of Xanthocnemis zealandica. Approximately 25 acts occur, a number comparable to the maximum repertoire sizes recorded for vertebrates and social insects, notwithstanding the absence of displays associated with reproductive activity in this insect larva. The behaviours of the territorial larvae of this species are compared with ethograms prepared from less extensive observations on the nonterritorial larvae of Ischnura aurora and Austrolestes colensonis. Both coenagrionid species used similar motor patterns in their displays; the lestid display involved a different motor pattern.
Larval growth of Lestes disjunctus was completed in 70 d in the field, whereas in Coenagrion resolutum it took 10–22 months. This was not simply the result of occupying warmer microhabitats, because L. disjunctus larvae grew faster than C. resolutum at all constant temperatures between 10 and 25 °C in the laboratory. Multiple regression analysis showed that growth rates of both species were positively related to temperature and negatively related to larval size and to the square of temperature. The latter term is necessary to describe the decrease in growth rate at high temperatures. The equation predicted that the growth rate of L. disjunctus reached a maximum at 28.8 °C, whereas that of C. resolutum decreased above 22.4 °C. Small and medium-sized larvae of L. disjunctus ate more prey (Daphnia magna) in 15 min than C. resolutum at all prey densities. In 15-min experiments the attack coefficient for small L. disjunctus larvae was significantly larger than for small C. resolutum larvae and handling time for medium larvae was shorter. Other comparisons had large associated sampling errors, but the trends were the same. These differences may be associated with the relatively longer labia of L. disjunctus and its ability to change hunting methods from ambush to active search.