Demise and rise : the biogeography and taxonomy of the Odonata of tropical Africa Dijkstra, K.D.B. Citation Dijkstra, K. D. B. (2007, May 16). Demise and rise : the biogeography and taxonomy of the Odonata of tropical Africa. Retrieved from https://hdl.handle.net/1887/11969 Version: Corrected Publisher’s Version Licence agreement concerning inclusion of doctoral thesis in License: the Institutional Repository of the University of Leiden Downloaded from: https://hdl.handle.net/1887/11969 Note: To cite this publication please use the final published version (if applicable). Demise and rise the biogeography and taxonomy of the Odonata of tropical Africa Klaas-Douwe B. Dijkstra Dijkstra, K.-D.B. 2007 Demise and rise: the biogeography and taxonomy of the Odonata of tropical Africa PhD Th esis, Leiden University Front cover photos: Gomphidia gamblesi (above) and Kintampo Falls, Ghana, by K.-D. B. Dijkstra Back cover photo: the author in Ankasa, Ghana, by Eric F. Th omassen Lay-out: Guido O. Keijl Printed by Finesse Druk, Heerhugowaard Demise and rise the biogeography and taxonomy of the Odonata of tropical Africa Proefschrift ter verkrijging van de graad van Doctor aan de Universiteit Leiden, op gezag van de Rector Magnifi cus prof. mr. P.F. van der Heijden, volgens besluit van het College voor Promoties, te verdedigen op woensdag 16 mei 2007, klokke 13.45 uur. door Klaas-Douwe Benediktus Dijkstra geboren te Kampen in 1975 Promotiecommissie Promotor: Prof. dr. E. Gittenberger (Nationaal Natuurhistorisch Museum Naturalis; Universiteit Leiden) Co-promotor: Dr. V. Clausnitzer (Philipps-Universität Marburg, Duitsland) Referent: Dr. M.L. May (Rutgers University, New Brunswick, Verenigde Staten) Overige leden: Prof. dr. J.J.M. van Alphen (Universiteit Leiden) Prof. dr. P.M. Brakefi eld (Universiteit Leiden) Prof. dr. P.J.J. Hooykaas (Universiteit Leiden) Dr. M. Schilthuizen (Nationaal Natuurhistorisch Museum Naturalis) Water No Get Enemy Toba fe lo weh omi lo malo If you wan go wash, water you go use Toba fe sobeh omi lo malo If you wan cook soup, water you go use To ri ba ngbona omi lero lero If your head be hot, water it cool am Tomo ba ngagda omi lo malo If your child dey grow, water you go use If water kill your child, water you go use Tobi ba bwi nao homi lo malo Ko sohun tole se ko ma lomi o Nothing without water Water, it no get enemy Fela Kuti (1975) 30 June 2006 Kintampo, Ghana for beauty CONTENTS chapter 1 A review of the taxonomy of African Odonata: fi nding ways to better identifi cation and biogeographic insight .....................................................................9 Cimbebasia 18: 191-206, 2003 chapter 2 Tropical African Platycnemis damselfl ies (Odonata: Platycnemididae) and the biogeographical signifi cance of a new species from Pemba Island, Tanzania ............27 Systematics & Biodiversity 5(2), 2007, in press chapter 3 Th e Pseudagrion split: molecular phylogeny confi rms the morphological and ecological dichotomy of Africa’s most diverse genus of Odonata (Coenagrionidae) .41 International Journal of Odonatology 10: 31-41, 2007 chapter 4 Gone with the wind: westward dispersal across the Indian Ocean and island speciation in Hemicordulia dragonfl ies (Odonata: Corduliidae) ................................55 Zootaxa 1438: 27-48, 2007 chapter 5 Two new relict Syncordulia species found during museum and fi eld studies of threatened dragonfl ies in the Cape Floristic Region (Odonata: Corduliidae) ...........79 Zootaxa 2007, in press chapter 6 Th e Atoconeura problem revisited: taxonomy, phylogeny and biogeography of a dragonfl y genus in the highlands of Africa (Odonata, Libellulidae) ......................97 Tijdschrift voor Entomologie 149: 121-144, 2006 chapter 7 Taxonomy and biogeography of Porpax, a dragonfl y genus centred in the Congo Basin (Odonata, Libellulidae) .....................................................................123 Tijdschrift voor Entomologie 149: 71-88, 2006 chapter 8 Demise and rise: the biogeography and taxonomy of the Odonata of tropical Africa ......................................................................................................................199 samenvatting Vergaan en verrijzen: de biogeografi e en taxonomie van de Odonata van tropisch Afrika ........................................................................................................189 a word of gratitude ..............................................................................................................................199 curriculum vitae ................................................................................................................................200 publications ................................................................................................................................202 8 chapter 1 A review of the taxonomy of African Odonata: fi nding ways to better identifi cation and biogeographic insight Cimbebasia 18: 191-206, 2003 Klaas-Douwe B. Dijkstra 10 chapter 1 - review of African Odonata taxonomy A review of the taxonomy of African Odonata - finding ways to better identification and biogeographic insight Klaas-Douwe B. Dijkstra Gortestraat 11, NL-2311 MS Leiden, The Netherlands: e-mail: [email protected] The taxonomy of the approximately 850 species of sub-Saharan African Odonata is relatively well-known, probably due to the impoverished nature of the fauna as compared to that of other tropical regions. The need for revisions, study of higher classification, comprehension of (often clinal, environmentally induced) variability and knowledge of larvae, phylogeny and biogeogra- phy are stressed. Taxonomic priorities are discussed for each family. Supportive activities include the production of identification manuals for a broader public, the accumulation of supplemen- tary material and the conservation of existing collections. A list of genera with estimated num- bers of species, taxonomic status and references is provided, as well as a list of important regional works. INTRODUCTION biogeography. The first priority for the taxonomy of African Odonata is, therefore, revisions. Although not as well studied as the Holarctic and Australasian Region’s faunas, knowledge of the REVISIONS & REVIEWS taxonomy of African Odonata is well ahead of those of the Oriental and Neotropic Regions. The It is hardly an exaggeration to say that all genera main reason is that the African fauna is relatively require at least some study. Elliot Pinhey tackled impoverished, harbouring only about 60% of the many problems, as can be seen from the nume- number of species found in each of the two other rous referrals to his work in the Appendix. Due to tropical regions. The writer here defines the Afri- their size, large genera generally have the greatest can fauna as that occurring south of the Sahara, problems. Pseudagrion is by far the largest genus with the inclusion of the Indian Ocean islands with almost a hundred species. Phyllomacromia and (Comoros, Madagascar, Mascarenes and Sey- Trithemis both number around forty species, Chloro- chelles). It numbers around 850 species, placed in cypha, Paragomphus and Orthetrum nearly thirty, and 125 genera. The former figure is about 15% of the Ceriagrion, Platycnemis, Gynacantha, Notogomphus and World total. Around 95% of species are not found Phyllogomphus about twenty. Other larger genera beyond the region. Despite the fact that the Afri- are Lestes, Nesolestes, Agriocnemis, Elattoneura and can Odonata are relatively well-known, numerous Zygonyx. Subtraction of the well-studied genera taxonomic issues remain uncertain, or at least from the largest genera creates an idea of revision unsettlingly hazy. Elucidating the relationships be- priority (vide Appendix). Besides full-scale revisions, tween species, and finding ways to distinguish many smaller issues of suspected synonyms, no- them, is perhaps the primary challenge of African menclature, mix-ups, identification problems and odonatology. This knowledge is required as a fun- species limits require attention. Some of the larger dament for future research, particularly for the al- genera that appear fairly well sorted could be re- most untouched field of African odonate viewed to straighten out the identification of the Proceedings of the 1st PHAON Meeting presented at the 2nd WDA International Symposium of Odonatology, Gällivare, Sweden, 26th July 2001. 11 demise and rise: tropical African dragonfl ies species. The problems are reviewed for each family backgrounds of them might be. Examples are the below. forms of Palpopleura lucia (Drury, 1773) (e.g. O’Neill & Paulson 2001) and Eleuthemis buettikoferi Ris, HIGHER CLASSIFICATION 1910 (Lempert 1988) and the subspecies of Atoco- neura biordinata (Karsch, 1899) (Longfield 1953). Next to taxonomic work with a species-group (most often genus) approach, there are numerous LARVAE problems in higher classification that require to be addressed. These are often relative to the fauna of Most of what has been published on African Odo- the Oriental Region, with which the African fauna nata (and most of what is written in this review) is has most in common. It is a lucky coincidence that about adults, but perhaps the greatest taxonomic when Elliot Pinhey’s ‘trailblazer’ Frederic Fraser frontier lies in the field of the larval stages. Al- began studying the African fauna intensively in though their study has (logically) lapsed behind, the 1940s, he had about twenty years of experi- the larvae or exuviae of many genera have been ence of the oriental fauna.