ArticlePDF Available

Sixty new dragonfly and damselfly species from Africa (Odonata)

Authors:
Kd Dijkstra at Naturalis Biodiversity Center
Kd Dijkstra
Jens Kipping at BioCart Ökologische Gutachten - Environmental Consultants
Jens Kipping
  • BioCart Ökologische Gutachten - Environmental Consultants
Nicolas Mézière at Ministère de l'Éducation nationale
Nicolas Mézière
Download full-text PDF
Read full-text
Download citation

Abstract

Man knows just one fifth of the nine million species of animal, plant, fungus and protist thought to inhabit our planet. Dragonflies and dam. selflies are regarded as well-known, however. Nevertheless we describe 60 new species, the most to be named at once in 130 years, adding one to every twelve species known in Africa. Each species is colourful and can often be recognised even from a photograph, showing that not all unknown life is indistinct and concealed. The species' beauty and sensitivity can raise awareness for the densest and most threatened biodiversity: freshwater covers less than one percent of Earth's surface, but harbours ten percent of animal species, of which a third may be at risk of extinction. Most of them, like dragonflies, are insects. They are popular indicators of habitat value and quality, but without a name cannot be added to the IUCN Red List. As habitats are rapidly disappearing, more exploratory and descriptive research is needed, support for which has waned. Nature, natural historians and the archives of life they build together are all under threat: our 60 new species are therefore as much an act of desperation as urgency.
Content uploaded by Jens Kipping
Author content
All content in this area was uploaded by Jens Kipping on Dec 05, 2015
Content may be subject to copyright.
Sixty new dragony and damsely species from Africa 447
Odonatologica 44(4) 2015: 447-678
1st December 2015
Sixty new dragony and damsely species
from Africa (Odonata)
Klaas-Douwe B. Dijkstra1, Jens Kipping2 & Nicolas Mézière3
1 Department of Conservation Ecology and Entomology,
University of Stellenbosch, South Africa & Naturalis Biodiversity Center,
P.O. Box 9517, 2300 RA Leiden, e Netherlands; <kd.dijkstra@naturalis.nl>
2 BioCart Environmental Consulting, Albrecht-Dürer-Weg 8,
04425 Taucha/Leipzig, Germany; <biocartkipping@email.de>
3 7 Avenue du Bois Chaudat, 97310 Kourou, French Guyana; <nicomez@free.fr>
Received and accepted 8th October 2015
Taxonomic abstract. e following new species of Odonata are described from Africa:
Umma gumma, Africocypha varicolor, Chlorocypha aurora, Chlorocypha ammea, Chloro-
cypha granata, Chlorocypha maxima, Pentaphlebia mangana, Allocnemis vicki, Elattoneura
aurifex, Elattoneura lapidaria, Elattoneura tarbotonorum, Aciagrion bapepe, Africallagma
quingentum, Agriocnemis canuango, Agriocnemis toto, Ceriagrion banditum, Ceriagrion
junceum, Ceriagrion obfuscans, Pseudagrion aureolum, Pseudagrion dactylidium, Pseud agrion
munte, Pseudagrion pacale, Pseudagrion sarepi, Pseudagrion tanganyicum, Anax gladiator,
Gynacantha congolica, Gynacantha pupillata, Lestinogomphus calcaratus, Lestinogomphus
nefrens, Lestinogomphus obtusus, Lestinogomphus venustus, Notogomphus bosumbuli, Noto-
gomphus cobyae, Notogomphus gorilla, Notogomphus intermedius, Notogomphus kimpavita,
Onychogomphus undecim, Paragomphus cammaertsi, Paragomphus clausnitzerorum, Para-
gomphus darwalli, Paragomphus dispar, Paragomphus lemperti, Phyllogomphus bongorum,
Tragogomphus grogona, Eleuthemis eogaster, Eleuthemis libera, Eleuthemis umbrina, Mal-
gassophlebia andzaba, Neodythemis infra, Neodythemis katanga, Orthetrum agaricum, Orthe-
trum kafwi, Orthetrum lusinga, Orthetrum umbratum, Porpax mezierei, Trithemis hinnula,
Trithemis legrandi, Urothemis venata, Zygonyx annika, Zygonyx denticulatus and Zygonyx
dionyx. e taxonomy of these genera and species-groups and complexes are also discussed:
Chlorocypha, including the diagnosis of C. dahli, C. ghesquierei and C. victoriae; the pauli-
group of Allocnemis; the glauca-group and vrijdaghi-complex of Elattoneura; the suave-com-
plex of Ceriagrion, including the diagnosis of C. mourae, C. sakejii and C. suave; the varians-
group of Ceriagrion, including the diagnosis and rejected synonymy of C. platystigma with
C. varians; the speratus-group of Anax, including the diagnosis and rejected synonymy of
A.rutherfordi with A. speratus; the bullata-group of Gynacantha, including the diagnosis and
rejected synonymy of G. victoriae with G. bullata; Lestinogomphus, including the diagnosis of
L. matilei and new synonymy of L. (formerly Microgomphus) bivittatus with Mastigogomphus
(formerly Neurogomphus) chapini; Notogomphus, including the new synonymies of N. buto-
loensis with N. leroyi, of N. anaci and N. verschuereni with N. spinosus, and of N. meruensis
K.-D.B. Dijkstra, J. Kipping & N. Mézière448
Odonatologica 44(4) 2015: 447-678
with N. kilimandjaricus; the supinus-group of Onychogomphus; Paragomphus, especially the
cognatus-group, including the diagnoses and new synonymies of P. bredoi and P. xanthus
with P. serrulatus, and of P. interruptus with P. machadoi, and the diagnosis of P. maynei; Eleu-
themis, including the diagnosis and rejected synonymy of E. quadrigutta with E. buettikoferi;
the saegeri-group of Orthetrum; the basitincta- and longistyla-groups of Trithemis, including
the new synonymies of the genera Anectothemis, Congothemis, Lokithemis and Porpacithemis
with Trithemis, and of T. trithemoides with T. apicalis; and the avicosta-complex of Zygonyx.
Man knows just one h of the nine million species of animal, plant,
fungus and protist thought to inhabit our planet. Dragonies and dam-
selies are regarded as well-known, however. Nevertheless we describe 60
new species, the most to be named at once in 130 years, adding one to eve-
ry twelve species known in Africa. Each species is colourful and can oen
be recognised even from a photograph, showing that not all unknown life
is indistinct and concealed. e species’ beauty and sensitivity can raise
awareness for the densest and most threatened biodiversity: freshwater
covers less than one percent of Earth’s surface, but harbours ten percent
of animal species, of which a third may be at risk of extinction. Most of
them, like dragonies, are insects. ey are popular indicators of habitat
value and quality, but without a name cannot be added to the IUCN Red
List. As habitats are rapidly disappearing, more exploratory and descrip-
tive research is needed, support for which has waned. Nature, natural his-
torians and the archives of life they build together are all under threat:
our 60 new species are therefore as much an act of desperation as urgency.

Supplementary resource (1)

GBIF Occurrence Download
Data
January 2016
occdownload gbif.org
... The information provided by surveys becomes very useful in assessing the conservation status of species, which leads to a more targeted conservation effort. The inclusion of more African Odonata in the global IUCN Red List (Dijkstra et al. 2009(Dijkstra et al. , 2011Clausnitzer and Dijkstra 2018;Clausnitzer 2020) after sufficient information on species level had been made available, based on a series of revisions and new species records (e.g., Dijkstra, Kipping, and Mézière 2015) published as part of continuously targeted expeditions in Africa, is a practical example of the success of targeted surveys and expeditions in Africa. ...
... Hence, conservation efforts on the species can only be achieved through the umbrella approach until additional information about their ecology and distribution are obtained. Explorative efforts to rediscover poorly known or rare populations should be intensified particularly in West Africa (Dijkstra et al. 2015). This implies that field expeditions by West African natural scientists should be intensified. ...
New records of some aquatic insect species in Nigeria: a need to intensify field expeditions in West Africa
Article
  • Jul 2024
  • AQUAT INSECT
A number of expeditions to eight freshwater systems within and around protected areas in the southern region of Nigeria were carried out between 2019 and 2022, with a view to re-discovering relict, rare, and poorly known aquatic invertebrate species. A total of 167 species from 10 orders were collected. Four species were recorded for the first time from Nigeria: Chlorocypha luminosa Karsch, 1893 and C. glauca Selys, 1879 (Odonata: Chlorocyphidae), Ephoron savignyi Williamson, 1802 (Ephemeroptera: Polymitarcyidae), and Cylindrostethus quadrivittatus Bergroth, 1916 (Hemiptera: Gerridae). Findings from this study also revealed that the distribution ranges of Elattoneura girardi Legrand, 1980 (Odonata: Platycnemididae) and Eurymetropsiella schoutedeni Poisson, 1950 (Hemiptera: Gerridae) extend to the southeastern region of Nigeria from the southwestern and northeastern regions, respectively. Rare and/or endemic species of Gerridae (Hemiptera) such as Eurymetra pauliani Poisson, 1941, Eurymetropsiella schoutedeni and Eurymetropsis carayoni Poisson, 1948 were also recorded in this study after several decades of paucity of information on their occurrence. The implications of aquatic invertebrate taxonomy and systematics on conservation in West Africa are discussed, and recommendations are provided.
View
... Implicitly, almost all "turbo-taxonomy" studies have embraced such faunistic sampling without much discussion (e.g. Fernandez-Triana et al., 2014, Fernandez-Triana, 2022Marsh et al., 2013;Riedel et al., 2013a, b;Riedel and Narakusumo, 2019, but see Dijkstra et al., 2015), presumably because it goes against the conventional recommendation that taxonomic revisions should cover all known species/specimens of a clade at a geographic scale appropriate for the species included in the clade. Such "completism" is desirable, but also contributes to taxonomic chauvinism because it cannot be applied to dark taxa. ...
"Dark taxonomy": A new protocol for overcoming the taxonomic impediments for dark taxa and broadening the taxon base for biodiversity assessment
Article
Full-text available
  • Feb 2025
  • CLADISTICS
We are entering the sixth mass extinction with little data for “dark taxa”, although they comprise most species. Much of the neglect is due to the fact that conventional taxonomic methods struggle with handling thousands of specimens belonging to hundreds of species. We thus here propose a new strategy that we call “dark taxonomy”. It addresses (i) taxonomic impediments, (ii) the lack of biodiversity baselines and (iii) the low impact of revisionary research. Taxonomic impediments are reduced by carrying out revisions at small geographic scales to keep the number of specimens low. The risk of taxonomic error is reduced by delimiting species based on two types of data. We furthermore show that dark taxonomy can yield important biodiversity baseline data by using samples obtained with biomonitoring traps. Lastly, we argue that the impact of revisionary research can be improved by publishing two papers addressing different readerships. The principles of dark taxonomy are illustrated by our taxonomic treatment of Singapore's fungus gnats (Mycetophilidae) based only on Malaise trap samples. We show that a first batch of specimens ( N = 1454) contains 120 species, of which 115 are new to science, thus reducing taxonomic impediments by increasing the number of described Oriental species by 25%. Species delimitation started with using DNA barcodes to estimate the number of Molecular Operational Taxonomic Units (MOTUs) before “LIT” (Large‐scale Integrative Taxonomy) was used to obtain the species boundaries for the 120 species by integrating morphological and molecular data. To test the taxonomic completeness of the revision, we next analysed a second batch of 1493 specimens and found that >97% belonged to the 120 species delimited based on the first batch. Indeed, the second batch only contained 18 new and rare MOTUs, i.e. our study suggests that a single revision can simultaneously yield the names for all important species and relevant biodiversity baseline data. Overall, we believe that “dark taxonomy” can quickly ready a large unknown taxon for biomonitoring.
View
... bispina Fraser, 1958; M. aequatoris Legrand, 1979;M. westfalli Legrand, 1986;and M. andzaba (Dijkstra & Mézière, 2015) and the subspecies M. bispina nigeriae Pinhey, 1961, were described from mainland Africa, as well as M. mediodentata Legrand, 2001, from Madagascar. Dijkstra (2007 considered M. aequatoris and M. bispi na nigeriae as well as Allorhizucha longistipes Pinhey, 1964, synonyms of M. bispina, leaving only three valid mainland African species. ...
View
... The first key principle of "dark taxonomy" is embracing "faunistic sampling", i.e., the geographic scope of a taxonomic revision is determined by the number of specimens and species that can be reasonably handled and not by the putative range of species. Implicitly, almost all "turbo-taxonomy" studies have embraced such sampling without much discussion (e.g., Fernandez-Triana et al. 2014Marsh et al., 2013;Riedel et al., 2013a,b;Riedel and Narakusumo 2019, but see Dijkstra et al. 2015). The lack of open discussion is presumably because it goes against the traditional recommendation that taxonomic revisions should cover all known species/specimens of a clade at a geographic scale appropriate for the species included in the clade. ...
"Dark taxonomy": a new protocol for overcoming the taxonomic impediments for dark taxa and broadening the taxon base for biodiversity assessment
Preprint
Full-text available
  • Sep 2023
We are entering the 6th mass extinction event on the planet with scarcely any data for "dark taxa" that comprise most animal species. These taxa have been neglected, because conventional taxonomic methods are not well-equipped to process tens of thousands of specimens belonging to thousands of species. We here test a new protocol for tackling the data deficiency problem for dark taxa by simultaneously addressing (1) taxonomic impediments, (2) lack of biodiversity baseline data, (3) and low impact of revisionary research. We propose to overcome the taxonomic impediments by carrying out revisions at small geographic scales to keep the number of specimens and species manageable. To lower the risk of taxonomic error, species delimitation and description are based on multiple data sources ("integrative taxonomy"). Secondly, we propose to create baseline data for dark taxa by initially revising them only based on fresh specimens obtained with the same sampling methods that are also used for biodiversity monitoring (e.g., Malaise traps for flying insects). This ensures that the revision constitutes a biodiversity baseline and the species most relevant for biomonitoring are revised first. Thirdly, we propose to improve the impact of taxonomic revisions by publishing two papers addressing different readerships (general and specialists). We illustrate our proposals by carrying out a taxonomic revision of the fungus gnats (Diptera: Mycetophilidae) of Singapore based on specimens obtained with Malaise traps placed at 107 sites across different habitats. We show that a first batch of specimens (N=1,456) contains 120 species, of which 115 are new to science and described in a separate taxonomic monograph. Species delimitation started with obtaining NGS barcodes (313-bp) that were used for estimating the number of MOTUs. This revealed 116-129 MOTUs depending on whether PTP, ABGD (P=0.001-0.060), or objective clustering (2-5%) was used. Afterwards, we applied the "LIT" process (Large-scale Integrative Taxonomy) which optimizes the integration of morphological and molecular data by using the molecular data to identify which specimens should be studied morphologically. Applying LIT to 3% MOTUs revealed a match ratio of 91% between the MOTUs and putative morphospecies, but an even higher match ratio of 97% was found for MOTUs obtained with 5% Objective Clustering and ABGD (P=0.060). Using these parameters, only two MOTUs had to be fused for perfect congruence between molecular and morphological data for the 120 species. To test the completeness of our monograph, we then analyzed a second batch of 1,567 specimens. Here, all clustering algorithms and settings revealed only 20 additional MOTUs. This means that the first batch of specimens already covered 85% of the diversity, although 22% and 10% of the species are singletons or doubletons, respectively. Even more remarkable is that >97% of all specimens in the second batch belonged to species described based on the first batch. The study thus demonstrates that revisions of dark taxa at moderate geographic scales with a moderate number of specimens from standardized traps can move dark taxa from being unavailable for biomonitoring to being well suited. Incidentally, the study also increases the number of described species of Mycetophilidae in the entire Oriental Region by >25% indicating the size of taxonomic impediments for dark taxa in Southeast Asia.
View
... Worryingly, many of these leaststudied regions not only harbor some of the most diverse and threatened odonate assemblages on the planet (Clausnitzer et al. 2012), but also suffer the fastest habitat degradation from land use and climate change (e.g. Southeast Asia, Africa) (Barlow et al. 2018;Dijkstra et al. 2015). Therefore, we do have a major weakness in our intent to understand and manage the global diversity of odonates. ...
Bridging people and nature through Odonata
Chapter
  • Nov 2022
This research-level text documents the latest advances in odonate biology and relates these to a broader ecological and evolutionary research agenda. Despite being one of the smallest insect orders, dragonflies offer a number of advantages for both laboratory and field studies. In fact, they continue to make a crucial contribution to the advancement of our broader understanding of insect ecology and evolution. This new edition provides a critical summary of the major advances in these fields. Contributions from many of the leading researchers in dragonfly biology offer new perspectives and paradigms as well as additional unpublished data. The editors have carefully assembled a mix of theoretical and applied chapters (including those addressing conservation and monitoring) as well as a balance of emerging (e.g. molecular evolution) and established research topics, providing suggestions for future study in each case. This accessible text is not about dragonflies per se, but rather an essential source of knowledge that describes how different sets of evolutionary and ecological principles/ideas have been tested on a particular taxon. This second edition of Dragonflies and Damselflies is suitable for graduate students and researchers in entomology, evolutionary biology, population and behavioral ecology, community ecology, and conservation biology. It will be of particular interest and use to those working on insects and an indispensable reference text for odonate biologists.
View
Molecular characterization and genetic divergence among the species of families Aeshnidae and Macromiidae (Odonata: Anisoptera) based on mitochondrial COI gene
Article
Full-text available
  • Mar 2025
DNA barcoding based on 23 COI sequences referable to 16 sequences of 10 species (Anaciaeschna jaspidea, Anax ephippiger, Anax guttatus, Anax immaculifrons, Anax indicus, Anax nigrofasciatus nigrolineatus, Anax parthenope, Gynacantha bainbriggei, Gynacantha bayadera, Gynacantha subinterrupta) of family Aeshnidae and 7 sequences of 4 species (Epophthalmia vittata, Macromia ellisoni, Macromia flavicincta, Macromia moorei) of family Macromiidae have been done. Tramea carolina species of the family Libellulidae is considered as out-group. Genetic divergence among the species of both the families based on variable, parsimony informative, conserved sites, nucleotide base composition of COI gene fragment and transversion/transistional bias have been calculated. COI gene sequences of three species Anax immaculifrons, Anax indicus and Gynacantha bainbriggei of family Aeshnidae and four species of Epophthalmia vittata, Macromia ellisoni, Macromia flavicincta and Macromia moorei of family Macromiidae have been submitted for the first time to NCBI, while sequences of 5 species Anax ephippiger, Anax guttatus, Anax nigrofasciatus nigrolineatus, Anax parthenope and Gynacantha subinterrupta matched with corresponding sequences of the species of family Aeshnidae have been submitted for the first time from India.
View
Description of the Larva of Argiocnemis rubescens rubeola Selys, 1877 (Odonata, Coenagrionidae, Agriocnemidinae) from Thailand with Systematic Notes on the Subfamily Agriocnemidinae
Article
  • May 2024
The final stadium larvae of Argiocnemis rubescens rubeola are described in the present study. The adult–larva association along with the taxonomic remarks on the subfamily Agriocneminae, is established by employing the cytochrome oxidase subunit I gene (COI). The larva is characterized by the ovate shape of the caudal lamella, the lack of a band on the caudal lamella, a rounded postocular lobe outline, three pairs of premental setae, and four palpal setae. Taxonomic notes suggest the COI data consistently aligns with both larval and adult characteristics, offering a comprehensive perspective.
View
DNA barcoding and molecular phylogeny of genus Pseudagrion (Odonata: Coenagrionidae) based on mitochondrial COI, ND1 and 16S rRNA genes
Article
Full-text available
  • May 2024
  • Int J Trop Insect Sci
Genus Pseudagrion is one of the largest genera of order Odonata and 14 species of this genus have been reported in India. The current research aims to reconstruct the phylogenetic trees by amplifying the partial sequences of the following three mitochondrial genes: COI, ND1 and 16 S rRNA genes, to produce a DNA reference library and determine the evolutionary relationships among the species of genus Pseudagrion. Sequences of mitochondrial genes of eight species have been submitted to NCBI, out of which Pseudagrion decorum, P. hypermelas and P. indicum barcoded for the first time. Since no work has been reported to far for CUB in genus Pseudagrion, the MT-ND1 gene appears to be a good choice for analyzing codon usage pattern. Codon usage bias for mitochondrial genes is low because of high effective number of codons. The most frequent codons, favored A or T at the 3rd codon position which indicate the significant role of compositional constraint in codon usage bias. Genetic divergence based on COI and 16 S rRNA ranges from 2.4 to 15.8%. 12 S rRNA and 16 S rRNA genes show minor genetic diversity, whereas NDI and COI genes reveal higher genetic diversity because of high mutation rate. In phylogenetic tree, 34 Pseudagrion species get segregated into five major clades as: ((P. pruinosum + P. spencei + P. hypermelas + P. rubriceps) + P. microcephalum) + (P. decorum + P. indicum + P. australasiae) + ((P. aureolum) + P. massaicum + P. tanganyicum + P. hamoni + P. pacale + P. niloticum + P. acacia) + (P. ignifer + P. pilidorsum + P. pruinosum) + ((P. dactylidium) + P. bicoerulans + P. munte + P. sarepi + P. kersteni) that depicts their evolutionary relationships.
View
Freshwater macroinvertebrates along the Nigeria-Cameroon border enhance the conservation value of the lower Guinea forest biodiversity hotspot
Article
  • Mar 2024
  • J ENVIRON MANAGE
In Nigeria and tropical Africa, the bias towards the protection of natural sites on account of terrestrial biodiversity or their resultant ecosystem services has led to a considerably low number of freshwater conservation studies in the region. To close this gap, six lotic freshwater systems in three different natural sites (Obudu Mountains, Agbokim Waterfalls, and Cross River National Park) along the Cameroon border of Nigeria were evaluated for the biodiversity of macroinvertebrates and some aspects of water quality in December 2021 and March 2022. Among other macroinvertebrate-related indices, the community conservation index (CCI), which considers the presence of threatened, rare, and notable species, was used for the conservation evaluation of the freshwater systems. The Ephemeroptera-Plecoptera-Trichoptera (EPT) species were notably rich at the sites, with over 12 species each at the Afundu Stream, Kwa River, Cataract Stream, and Agbokim River. All of the sites had a very high conservation value, according to the CCI, which was greater than 20. Some Obudu sites (Grotto and Becheve streams), despite having fewer EPT taxa, had higher CCI than the Kwa or Agbokim River. This was due to the presence of the endangered damselflies (Africocypha centripunctata and Allocnemis vicki) at the Obudu sites and a rare mayfly species in Nigeria (Tricorythus tinctus), which was reported at Agbokim and Kwa Rivers. The presence of a notable damselfly genus (Pentaphlebia) at the Afundu stream of the Obudu Mountains also raises some hope that the critically endangered damselfly (Pentaphlebia gamblesi), whose adult was last seen in 1973, could still be alive at the site. Canonical Correspondence Analysis revealed that the proportion of scrapers had a strong association with high BOD and EC in Agbokim River, while the Becheve and Grotto streams which had high vegetal cover was positively associated with shredders. Also, increasing DO in Afundu Stream, Kwa River and Cataract Stream was positively associated with EPT richness. This study revealed that more freshwater ecological studies need to be conducted at isolated natural sites and protected areas in Africa. Findings from such studies have implications for preserving freshwater ecosystems of high conservation value from the various threats that characterize the Anthropocene.
View
Molecular Phylogeny of Holarctic Aeshnidae with a Focus on the West Palaearctic and Some Remarks on Its Genera Worldwide (Aeshnidae, Odonata)
Article
Full-text available
Aeshnidae Rambur, 1842 are impressive large insects distributed worldwide. Currently, over 500 species are recognized. Nevertheless, the phylogeny of this family is not completely understood. We applied molecular phylogenetic analysis using two popular phylogenetic markers, the mitochondrial COI gene fragment (barcoding sequence) and the nucleic ITS region, containing the ITS1, 5.8S rRNA, and ITS2 sequences. We used available and credible published sequences and 96 newly sequenced specimens. Our analysis involved all West Palaearctic species, all but one genera of the Holarctic Aeshnidae, and most genera worldwide, and is by far the largest molecular study of this family. The topology of all trees created with different algorithms and genes is in favour of the current taxonomic concept, with some remarkable outcomes. Aeshna Fabricius, 1775, was found to be diverged into several branches, especially with respect to the COI gene. Although it appeared not monophyletic in phylogenetic reconstructions based on the ITS region, the analysis of COI and joint analysis suggest its monophyly in the current taxonomical sense, with one notable exception. Aeshna isoceles (Müller, 1767) has fallen out of Aeshna in all analyses, so a new monophyletic genus, Isoaeschna gen. nov. is introduced for it. The genus Brachytron Evans, 1845 tightly clustered with Aeschnophlebia Selys, 1883, Epiaeschna Hagen in Selys, 1883, and Nasiaeschna Selys in Förster, 1900. Thus, we suggest subsuming these four genera under the priority name Brachytron. Tetracanthagyna Selys, 1883 clusters as expected with Brachytron in the ITS tree, but is an independent ancient clade of its own in all COI trees. The genus Polycanthagyna Fraser, 1933 syn. nov. is synonymised to Indaeschna Fraser, 1926. On the species level, we suggest that the American Aeshna septentrionalis Burmeister, 1839 be treated as a subspecies of A. caerulea (Ström, 1783), Aeshna caerulea septentrionalis. We synonymize Gynacantha hyalina Selys, 1882 with Gynacantha subinterrupta Rambur, 1842. Our analysis provides new insights on the tight relationships of the circumboreal species Aeshna juncea and A. subarctica and the intraspecies phylogeny of Aeshna juncea.
View
Genome-wide Evidence Reveals that African and Eurasian Golden Jackals Are Distinct Species
Article
Full-text available
  • Jul 2015
The golden jackal of Africa (Canis aureus) has long been considered a conspecific of jackals distributed throughout Eurasia, with the nearest source populations in the Middle East. However, two recent reports found that mitochondrial haplotypes of some African golden jackals aligned more closely to gray wolves (Canis lupus) [1, 2], which is surprising given the absence of gray wolves in Africa and the phenotypic divergence between the two species. Moreover, these results imply the existence of a previously unrecognized phylogenetically distinct species despite a long history of taxonomic work on African canids. To test the distinct-species hypothesis and understand the evolutionary history that would account for this puzzling result, we analyzed extensive genomic data including mitochondrial genome sequences, sequences from 20 autosomal loci (17 introns and 3 exon segments), microsatellite loci, X- and Y-linked zinc-finger protein gene (ZFX and ZFY) sequences, and whole-genome nuclear sequences in African and Eurasian golden jackals and gray wolves. Our results provide consistent and robust evidence that populations of golden jackals from Africa and Eurasia represent distinct monophyletic lineages separated for more than one million years, sufficient to merit formal recognition as different species: C. anthus (African golden wolf) and C. aureus (Eurasian golden jackal). Using morphologic data, we demonstrate a striking morphologic similarity between East African and Eurasian golden jackals, suggesting parallelism, which may have misled taxonomists and likely reflects uniquely intense interspecific competition in the East African carnivore guild. Our study shows how ecology can confound taxonomy if interspecific competition constrains size diversification. Copyright © 2015 Elsevier Ltd. All rights reserved.
View
Revision of the Sundaland species of the genus Dysphaea Selys, 1853 using molecular and morphological methods, with notes on allied species (Odonata: Euphaeidae)
Article
Full-text available
  • Apr 2015
  • ZOOTAXA
The Sundaland species of the genus Dysphaea were studied using molecular and morphological methods. Four species are recognized: D. dimidiata Selys, D. lugens Selys, D. ulu spec. nov. (holotype ♂, from Borneo, Sarawak, Miri division, Upper Baram, Sungai Pejelai, Ulu Moh, 24 viii 2014; deposited in RMNH) and D. vanida spec. nov. (holotype ♂, from Thailand, Ranong province, Khlong Nakha, Khlong Bang Man, 12–13 v 1999; deposited in RMNH). The four species are described and illustrated for both sexes, with keys provided. The type specimens of the four Dysphaea taxa named by E. de Selys Longchamps, i.e. dimidiata, limbata, semilimbata and lugens, were studied and their taxonomic status is dis-cussed. Lectotypes are designated for D. dimidiata and D. limbata. D. dimidiata is recorded from Palawan (the Philip-pines) for the first time. A molecular analysis using three markers (COI, 16S and 28S) is presented. This includes specimens of three Sundaland species of the genus (D. lugens missing) and two congeners from other regions (D. basi-tincta and D. gloriosa). Notes and photographs of the male holotype of D. walli Fraser (from Maymyo, Burma) are provided.
View
Review of Apanteles sensu stricto (Hymenoptera, Braconidae, Microgastrinae) from Area de Conservación Guanacaste, northwestern Costa Rica, with keys to all described species from Mesoamerica
Article
Full-text available
  • Feb 2014
More than half a million specimens of wild-caught Lepidoptera caterpillars have been reared for their parasitoids, identified, and DNA barcoded over a period of 34 years (and ongoing) from Area de Conservación de Guanacaste (ACG), northwestern Costa Rica. This provides the world's best location-based dataset for studying the taxonomy and host relationships of caterpillar parasitoids. Among Hymenoptera, Microgastrinae (Braconidae) is the most diverse and commonly encountered parasitoid subfamily, with many hundreds of species delineated to date, almost all undescribed. Here, we reassess the limits of the genus Apanteles sensu stricto, describe 186 new species from 3,200+ parasitized caterpillars of hundreds of ACG Lepidoptera species, and provide keys to all 205 described Apanteles from Mesoamerica - including 19 previously described species in addition to the new species. The Mesoamerican Apanteles are assigned to 32 species-groups, all but two of which are newly defined. Taxonomic keys are presented in two formats: traditional dichotomous print versions and links to electronic interactive versions (software Lucid 3.5). Numerous illustrations, computer-generated descriptions, distributional information, wasp biology, and DNA barcodes (where available) are presented for every species. All morphological terms are detailed and linked to the Hymenoptera Anatomy Ontology website. DNA barcodes (a standard fragment of the cytochrome c oxidase I (COI) mitochondrial gene), information on wasp biology (host records, solitary/gregariousness of wasp larvae), ratios of morphological features, and wasp microecological distributions were used to help clarify boundaries between morphologically cryptic species within species-complexes. Because of the high accuracy of host identification for about 80% of the wasp species studied, it was possible to analyze host relationships at a regional level. The ACG species of Apanteles attack mainly species of Hesperiidae, Elachistidae and Crambidae (Lepidoptera). About 90% of the wasp species with known host records seem to be monophagous or oligophagous at some level, parasitizing just one host family and commonly, just one species of caterpillar. Only 15 species (9%) parasitize species in more than one family, and some of these cases are likely to be found to be species complexes. We have used several information sources and techniques (traditional taxonomy, molecular, software-based, biology, and geography) to accelerate the process of finding and describing these new species in a hyperdiverse group such as Apanteles. The following new taxonomic and nomenclatural acts are proposed. Four species previously considered to be Apanteles are transferred to other microgastrine genera: Dolichogenidea hedyleptae (Muesebeck, 1958), comb. n., Dolichogenidea politiventris (Muesebeck, 1958), comb. n., Rhygoplitis sanctivincenti (Ashmead, 1900), comb. n., and Illidops scutellaris (Muesebeck, 1921), comb. rev. One European species that is a secondary homonym to a Mesoamerican species is removed from Apanteles and transferred to another genus: Iconella albinervis (Tobias, 1964), stat. rev. The name Apanteles albinervican Shenefelt, 1972, is an invalid replacement name for Apanteles albinervis (Cameron, 1904), stat. rev., and thus the later name is reinstated as valid. The following 186 species, all in Apanteles and all authored by Fernández-Triana, are described as species nova: adelinamoralesae, adrianachavarriae, adrianaguilarae, adrianguadamuzi, aichagirardae, aidalopezae, albanjimenezi, alejandromasisi, alejandromorai, minorcarmonai, alvarougaldei, federicomatarritai, anabellecordobae, rostermoragai, anamarencoae, anamartinesae, anapiedrae, anariasae, andreacalvoae, angelsolisi, arielopezi, bernardoespinozai, bernyapui, bettymarchenae, bienvenidachavarriae, calixtomoragai, carloscastilloi, carlosguadamuzi, eliethcantillanoae, carlosrodriguezi, carlosviquezi, carloszunigai, carolinacanoae, christianzunigai, cinthiabarrantesae, ciriloumanai, cristianalemani, cynthiacorderoae, deifiliadavilae, dickyui, didiguadamuzi, diegoalpizari, diegotorresi, diniamartinezae, duniagarciae, duvalierbricenoi, edgarjimenezi, edithlopezae, eduardoramirezi, edwinapui, eldarayae, erickduartei, esthercentenoae, eugeniaphilipsae, eulogiosequeira, felipechavarriai, felixcarmonai, fernandochavarriai, flormoralesae, franciscopizarroi, franciscoramirezi, freddyquesadai, freddysalazari, gabrielagutierrezae, garygibsoni, gerardobandoi, gerardosandovali, gladysrojasae, glenriverai, gloriasihezarae, guadaluperodriguezae, guillermopereirai, juanmatai, harryramirezi, hectorsolisi, humbertolopezi, inesolisae, irenecarrilloae, isaacbermudezi, isidrochaconi, isidrovillegasi, ivonnetranae, jairomoyai, javiercontrerasi, javierobandoi, javiersihezari, jesusbrenesi, jesusugaldei, jimmychevezi, johanvargasi, jorgecortesi, jorgehernandezi, josecalvoi, josecortesi, josediazi, josejaramilloi, josemonteroi, joseperezi, joserasi, juanapui, juancarrilloi, juangazoi, juanhernandezi, juanlopezi, juanvictori, juliodiazi, juniorlopezi, keineraragoni, laurahuberae, laurenmoralesae, leninguadamuzi, leonelgarayi, lilliammenae, lisabearssae, luciariosae, luisbrizuelai, luiscanalesi, luiscantillanoi, luisgarciai, luisgaritai, luishernandezi, luislopezi, luisvargasi, manuelarayai, manuelpereirai, manuelriosi, manuelzumbadoi, marcobustosi, marcogonzalezi, marcovenicioi, mariachavarriae mariaguevarae, marialuisariasae, mariamendezae, marianopereirai, mariatorrentesae, sigifredomarini, marisolarroyoae, marisolnavarroae, marvinmendozai, mauriciogurdiani, milenagutierrezae, monicachavarriae, oscarchavesi, osvaldoespinozai, pablotranai, pabloumanai, pablovasquezi, paulaixcamparijae, luzmariaromeroae, petronariosae, randallgarciai, randallmartinezi, raulacevedoi, raulsolorsanoi, wadyobandoi, ricardocaleroi, robertmontanoi, robertoespinozai, robertovargasi, rodrigogamezi, rogerblancoi, rolandoramosi, rolandovegai, ronaldcastroi, ronaldgutierrezi, ronaldmurilloi, ronaldnavarroi, ronaldquirosi, ronaldzunigai, rosibelelizondoae, ruthfrancoae, sergiocascantei, sergioriosi, tiboshartae, vannesabrenesae, minornavarroi, victorbarrantesi, waldymedinai, wilbertharayai, williamcamposi, yeissonchavesi, yilbertalvaradoi, yolandarojasae, hazelcambroneroae, zeneidabolanosae.
View
A New Species of River Dolphin from Brazil or: How Little Do We Know Our Biodiversity
Article
Full-text available
True river dolphins are some of the rarest and most endangered of all vertebrates. They comprise relict evolutionary lineages of high taxonomic distinctness and conservation value, but are afforded little protection. We report the discovery of a new species of a river dolphin from the Araguaia River basin of Brazil, the first such discovery in nearly 100 years. The species is diagnosable by a series of molecular and morphological characters and diverged from its Amazonian sister taxon 2.08 million years ago. The estimated time of divergence corresponds to the separation of the Araguaia-Tocantins basin from the Amazon basin. This discovery highlights the immensity of the deficit in our knowledge of Neotropical biodiversity, as well as vulnerability of biodiversity to anthropogenic actions in an increasingly threatened landscape. We anticipate that this study will provide an impetus for the taxonomic and conservation reanalysis of other taxa shared between the Araguaia and Amazon aquatic ecosystems, as well as stimulate historical biogeographical analyses of the two basins.
View
Redefining the damselfly families: A comprehensive molecular phylogeny of Zygoptera (Odonata)
Article
Full-text available
  • Aug 2013
  • SYST ENTOMOL
An extensive molecular phylogenetic reconstruction of the suborder Zygoptera of the Odonata is presented, based on mitochondrial (16S, COI) and nuclear (28S) data of 59% of the 310 genera recognized and all (suspected) families except the monotypic Hemiphlebiidae. A partial reclassification is proposed, incorporating morphological characters. Many traditional families are recovered as monophyletic, but reorganization of the superfamily Coenagrionoidea into three families is proposed: Isostictidae, Platycnemididae and Coenagrionidae. Archboldargia Lieftinck, Hylaeargia Lieftinck, Palaiargia Förster, Papuargia Lieftinck and Onychargia Selys are transferred from Coenagrionidae to Platycnemididae, and Leptocnemis Selys, Oreocnemis Pinhey and Thaumatagrion Lieftinck from Platycnemididae to Coenagrionidae. Each geographically well-defined clade of Platycnemididae is recognized as a subfamily, and thus Disparoneurinae (i.e. Old World ‘Protoneuridae’) is incorporated, Calicnemiinae is restricted, and Allocnemidinae (type genus: Allocnemis Selys) subfam.n., Idiocnemidinae (type genus: Idiocnemis Selys) subfam.n. and Onychargiinae (type genus: Onychargia Selys) subfam.n. and Coperini trib.n. (type genus: Copera Kirby) are described. Half of Coenagrionidae belongs to a well-supported clade incorporating Coenagrion Kirby and the potential subfamilies Agriocnemidinae, Ischnurinae and Pseudagrioninae. The remainder is less well defined, but includes the Pseudostigmatidae and New World Protoneuridae that, with Argiinae and Teinobasinae, may prove valid subfamilies with further evidence. Ninety-two per cent of the genera formerly included in the polyphyletic Amphipterygidae and Megapodagrionidae were studied. Pentaphlebiidae, Rimanellidae and Devadattidae fam.n. (type genus: Devadatta Kirby) are separated from Amphipterygidae, and Argiolestidae, Heteragrionidae, Hypolestidae, Philogeniidae, Philosinidae and Thaumatoneuridae from Megapodagrionidae. Eight further groups formerly placed in the latter are identified, but are retained as incertae sedis; the validity of Lestoideidae, Philogangidae and Pseudolestidae is confirmed. For some families (e.g. Calopterygidae, Chlorocyphidae) a further subdivision is possible; Protostictinae subfam.n. (type genus: Protosticta Selys) is introduced in Platystictidae. Numerous new combinations are proposed in the Supporting Information. Many long-established families lack strong morphological apomorphies. In particular, venation is incongruent with molecular results, stressing the need to review fossil Odonata taxonomy: once defined by the reduction of the anal vein, Protoneuridae dissolves completely into six clades from five families.
View
Global patterns of freshwater species diversity, threat and endemism
Article
Full-text available
  • Jan 2014
  • GLOBAL ECOL BIOGEOGR
Aim: Global-scale studies are required to identify broad-scale patterns in the distributions of species, to evaluate the processes that determine diversity and to determine how similar or different these patterns and processes are among different groups of freshwater species. Broad-scale patterns of spatial variation in species distribution are central to many fundamental questions in macroecology and conservation biology. We aimed to evaluate how congruent three commonly used metrics of diversity were among taxa for six groups of freshwater species. Location: Global. Methods: We compiled geographical range data on 7083 freshwater species of mammals, amphibians, reptiles, fishes, crabs and crayfish to evaluate how species richness, richness of threatened species and endemism are distributed across freshwater ecosystems. We evaluated how congruent these measures of diversity were among taxa at a global level for a grid cell size of just under 1°. Results: We showed that although the risk of extinction faced by freshwater decapods is quite similar to that of freshwater vertebrates, there is a distinct lack of spatial congruence in geographical range between different taxonomic groups at this spatial scale, and a lack of congruence among three commonly used metrics of biodiversity. The risk of extinction for freshwater species was consistently higher than for their terrestrial counterparts. Main conclusions: We demonstrate that broad-scale patterns of species richness, threatened-species richness and endemism lack congruence among the six freshwater taxonomic groups examined. Invertebrate species are seldom taken into account in conservation planning. Our study suggests that both the metric of biodiversity and the identity of the taxa on which conservation decisions are based require careful consideration. As geographical range information becomes available for further sets of species, further testing will be warranted into the extent to which geographical variation in the richness of these six freshwater groups reflects broader patterns of biodiversity in fresh water.
View
Rigorous approaches to species delimitation have significant implications for African crocodilian systematic and conservation
Article
Full-text available
Accurate species delimitation is a central assumption of biology that, in groups such as the Crocodylia, is often hindered by highly conserved morphology and frequent introgression. In Africa, crocodilian systematics has been hampered by complex regional biogeography and confounded taxonomic history. We used rigorous molecular and morphological species delimitation methods to test the hypothesis that the slender-snouted crocodile (Mecistops cataphractus) is composed of multiple species corresponding to the Congolian and Guinean biogeographic zones. Speciation probability was assessed by using 11 mitochondrial and nuclear genes, and cranial morphology for over 100 specimens, representing the full geographical extent of the species distribution. Molecular Bayesian and phylogenetic species delimitation showed unanimous support for two Mecistops species isolated to the Upper Guinean and Congo (including Lower Guinean) biomes that were supported by 13 cranial characters capable of unambiguously diagnosing each species. Fossil-calibrated phylogenetic reconstruction estimated that the species split ± 6.5-7.5 Ma, which is congruent with intraspecies divergence within the sympatric crocodile genus Osteolaemus and the formation of the Cameroon Volcanic Line. Our results underscore the necessity of comprehensive phylogeographic analyses within currently recognized taxa to detect cryptic species within the Crocodylia. We recommend that the community of crocodilian researchers reconsider the conceptualization of crocodilian species especially in the light of the conservation ramifications for this economically and ecologically important group.
View
Integrating multiple lines of evidence to better understand the evolutionary divergence of humpback dolphins along their entire distribution range: A new dolphin species in Australian waters?
Article
Full-text available
  • Dec 2013
  • MOL ECOL
The conservation of humpback dolphins, distributed in coastal waters of the Indo-West Pacific and eastern Atlantic Oceans, has been hindered by a lack of understanding about the number of species in the genus (Sousa) and their population structure. To address this issue, we present a combined analysis of genetic and morphologic data collected from beach-cast, remote-biopsied and museum specimens from throughout the known Sousa range. We extracted genetic sequence data from 235 samples from extant populations and explored the mitochondrial control region and four nuclear introns through phylogenetic, population-level and population aggregation frameworks. In addition, 180 cranial specimens from the same geographical regions allowed comparisons of 24 morphological characters through multivariate analyses. The genetic and morphological data showed significant and concordant patterns of geographical segregation, which are typical for the kind of demographic isolation displayed by species units, across the Sousa genus distribution range. Based on our combined genetic and morphological analyses, there is convincing evidence for at least four species within the genus (S. teuszii in the Atlantic off West Africa, S. plumbea in the central and western Indian Ocean, S. chinensis in the eastern Indian and West Pacific Oceans, and a new as-yet-unnamed species off northern Australia).
View
Global threats to human water security and river biodiversity
Article
  • Sep 2010
Protecting the worlds freshwater resources requires diagnosing threats over a broad range of scales, from global to local. Here we present the first worldwide synthesis to jointly consider human and biodiversity perspectives on water security using a spatial framework that quantifies multiple stressors and accounts for downstream impacts. We find that nearly 80% of the worlds population is exposed to high levels of threat to water security. Massive investment in water technology enables rich nations to offset high stressor levels without remedying their underlying causes, whereas less wealthy nations remain vulnerable. A similar lack of precautionary investment jeopardizes biodiversity, with habitats associated with 65% of continental discharge classified as moderately to highly threatened. The cumulative threat framework offers a tool for prioritizing policy and management responses to this crisis, and underscores the necessity of limiting threats at their source instead of through costly remediation of symptoms in order to assure global water security for both humans and freshwater biodiversity.
View
Freshwater Biodiversity and Aquatic Insect Diversification
Article
  • Oct 2013
  • ANNU REV ENTOMOL
Inland waters cover less than 1% of Earth's surface but harbor more than 6% of all insect species: Nearly 100,000 species from 12 orders spend one or more life stages in freshwater. Little is known about how this remarkable diversity arose, although allopatric speciation and ecological adaptation are thought to be primary mechanisms. Freshwater habitats are highly susceptible to environmental change and exhibit marked ecological gradients. Standing waters appear to harbor more dispersive species than running waters, but there is little understanding of how this fundamental ecological difference has affected diversification. In contrast to the lack of evolutionary studies, the ecology and habitat preferences of aquatic insects have been intensively studied, in part because of their widespread use as bioindicators. The combination of phylogenetics with the extensive ecological data provides a promising avenue for future research, making aquatic insects highly suitable models for the study of ecological diversification. Expected final online publication date for the Annual Review of Entomology Volume 59 is January 07, 2014. Please see http://www.annualreviews.org/catalog/pubdates.aspx for revised estimates.
View