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Zootaxa, Biogeography and Divergence Time Estimation of The Zootaxa 2216: 22–36 (2009) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2009 · Magnolia Press ISSN 1175-5334 (online edition) Biogeography and divergence time estimation of the relict Cape dragonfly genus Syncordulia: global significance and implications for conservation JESSICA L. WARE1,3,4, JOHN P. SIMAIKA2 & MICHAEL J. SAMWAYS2 1Department of Entomology, Rutgers University, New Brunswick, NJ, USA 2Department of Conservation Ecology and Entomology, Stellenbosch University, South Africa 3Division of Invertebrate Zoology, American Museum of Natural History, New York, USA 4Corresponding author. E-mail: [email protected] Abstract Syncordulia (Odonata: Anisoptera: Libelluloidea) inhabits mostly cool mountainous streams in the Cape Floristic Region of South Africa. It is found at low densities in geographically restricted areas. Syncordulia is endemic to South Africa and, until recently, only two species were known, S. venator (Barnard, 1933) and S. gracilis (Burmeister 1839), both considered Vulnerable by the World Conservation Union (IUCN). Two new species, S. serendipator Dijsktra, Samways & Simaika 2007 and S. legator Dijsktra, Samways & Simaika 2007, were described from previously unrecognized museum specimens and new field collections. Here we corroborate the validity of these two new species using multiple genes and propose intergeneric relationships within Syncordulia. Molecular data from two independent gene fragments (nuclear 28S and ribosomal and cytochrome oxidase subunit I mitochondrial data) were sequenced and/or downloaded from GenBank for 7 libelluloid families, including 12 Syncordulia specimens (2 Syncordulia gracilis, 4 S. serendipator, 2 S. legator and 4 S. venator). The lower libelluloid group GSI (sensu Ware et al. 2007), a diverse group of non– corduliine taxa, is strongly supported as monophyletic. Syncordulia is well supported by both methods of phylogenetic analyses as a monophyletic group deeply nested within the GSI clade. A DIVA biogeographical analysis suggests that the ancestor to the genus Syncordulia may have arisen consequent to the break–up of Gondwana (>120 Mya). Divergence time estimates suggest that Syncordulia diverged well after the breakup of Gondwana, approximately 60 million years ago (Mya), which coincides with the divergence of several Cape fynbos taxa, between 86 – 60 Mya. DIVA analyses suggest that the present distributions of Syncordulia may be the result of dispersal events. We relate these phylogenetic data to the historical biogeography of the genus and to the importance of conservation action. Key words: Syncordulia, Odonata, Biogeography, Divergence Times, Conservation Introduction Corduliid dragonflies, often known as “emeralds” for the metallic green body and eye color of many species, are members of the large superfamily, Libelluloidea. Previously considered to be one family (Rambur 1842; Hagen 1861; Kirby 1890; Selys-Longchamps 1892; Needham 1908; Martin 1914; Tillyard 1917 1928; St. Quentin 1939; Gloyd 1959; Lieftinck 1971; Lieftinck 1977; Carle 1982; Davies & Tobin 1985; Bridges 1994; Steinmann 1997) or divided into two or more families (Fraser 1957; Carle 1995; Bechly 1996; Lohmann, 1996a, b; Pfau, 2005), Corduliidae has not been found to be monophyletic in modern phylogenetic analyses (May 1995b; Misof et al. 2001; Carle & Kjer 2002; Ware et al. 2007; Bybee et al. 2008; Letsch unpubl.). ‘Corduliidae’ (sensu Fraser 1957) consists of at least three distinct clades, the Corduliinae, the Macromiinae, and a diverse group of non-corduliine taxa, informally called the “GSI” by Ware et al. (2007) after the subfamilies Gomphomacromiinae, Synthemistinae, and Idionychinae. Within the GSI are Australian endemics, as well as African, South American, European, and Indomalayan genera, and the presumably relict South African genus Syncordulia. 22 Accepted by R. Garrison: 15 Jul. 2009; published: 1 Sept. 2009 Although both GSI adults and larvae have been documented in the literature (Selys-Longchamps 1871; Geijskes 1970; Lieftinck 1960; Lieftinck 1971; Lieftinck 1977; May 1991; Theischinger & Watson 1978, 1984; Carle 1995; May 1995a, b; Carvalho et al. 2004; von Ellenrieder & Garrison 2005), the taxa have been infrequently used in modern phylogenetic studies (Carle 1995; Carle & Kjer 2002; Misof et al. 2001; Ware et al. 2007; Bybee et al. 2008, Letsch unpubl., Fleck et al. 2008). Of these studies, only Ware et al. (2007) incorporated Syncordulia in their phylogeny. The GSI stem forms a basal node in the Libelluloidea. Thus, the evolution of certain libelluloid apomorphies, such as reduction in the ovipositor for exophytic oviposition and modification of the distal sperm pump of the penis (Pfau 2005), cannot be fully evaluated without an understanding of intergeneric relationships within GSI. In addition, several members of the GSI are rare endemics, and phylogenetic hypotheses about this group may be useful in understanding the biogeography and conservation needs of vulnerable genera such as Syncordulia. First described by Selys-Longchamps in 1882, the genus Syncordulia comprises dragonflies endemic to South Africa and restricted almost entirely to the Cape Floristic Region (CFR). Inhabiting mostly cool mountainous streams in the CFR of South Africa, Syncordulia species generally occur in geographically- restricted areas and at low population densities. All known Syncordulia species occur in sympatry (Dijkstra et al., 2007) and are often found dwelling in the same stream. Until recently, only S. venator and S. gracilis were known, and the World Conservation Union (IUCN) (2001) considers both Vulnerable. Dijkstra et al., (2007) described two new species, S. serendipator and S. legator, from previously unrecognized museum specimens and from new field collections. Intrageneric relationships within Syncordulia have not yet been fully evaluated within a phylogenetic context. In addition, placement of Syncordulia within the GSI has been unresolved. Lieftinck (1960) suggested that Syncordulia was closely related to Oxygastra Selys 1870, Hesperocordulia Tillyard 1911, Lathrocordulia, Tillyard 1911 or Micromidia, Fraser 1959, but his study was pre-cladistic, and done without a formal analysis. Ware et al. (2007) included S. gracilis in their molecular analysis, but its position within the monophyletic GSI assemblage was unstable and had low branch support. Monophyly of the genus itself, although seemingly very likely based on several morphological characters including uniform wing vein patterns (shape of the anal loop, patterns of veins in the antenodal region, veins in the discoidal field, etc) and eye color, has not yet been supported by phylogenetic analysis. The validity of the two new Syncordulia species, and the monophyly of the genus are assessed in this study. In addition, we attempt to determine the placement of Syncordulia within the GSI. Using the resulting phylogenetic hypotheses, we estimate both divergence times and biogeographical history to better understand the distribution, ecology, and conservation requirements of this relict genus. Materials and methods Taxon selection. Although this study’s focus was on the relationships within Syncordulia, we included several other libelluloid taxa as outgroups (i.e., gomphomacromiine and idomacromiine taxa, sensu Davies and Tobin, 1985, Macromia, corduliine taxa and lower libelluloids, sensu Ware et al. 2007). For a better understanding of the relationships within GSI (sensu Ware et al. 2007), one would need several additional African taxa (such as Neophya, Idomacromia or Nesocordulia), and/or South American taxa (such as Lauromacromia, or Neocordulia) for which fresh material was unavailable to us (DNA extraction was attempted for each of these species but was unsuccessful). Future work on the GSI should attempt to include these taxa, although obtaining fresh specimens of these rare species will likely prove difficult. As outgroups we used Chlorogomphidae, Cordulegastridae and Neopetalia. Data collection. Specimens were collected in South Africa during the summer months of 2006 and 2007 (five S. serendipator, four S. legator and ten S. venator) and 2004 (four S. gracilis). DNA was extracted using a Qiagen DNEasy kit with a modified protocol as described by Ware et al. (2007). PCR amplification was SYNCORDULIA PHYLOGENY AND BIOGEOGRAPHY Zootaxa 2216 © 2009 Magnolia Press · 23 TABLE 1. Taxon samples from the present study. Species of Syncordulia GenBank Accession Collection information Numbers Date Sex Habitat Syncordulia gracilis COI: GQ329638 2004 Female In grasslands, no water D2/D3:None D7: None 16S: GQ329650 Syncordulia gracilis COI: GQ329637 2004 Female In grasslands, no water D2/D3: None D7: GQ329657 16S: None Syncordulia legator COI: GQ329636 18/10/2006 Female Open montane stream, D2/D3: GQ329640 riffles, glides, deposition D7: GQ329656 zones 16S: GQ329649 Syncordulia legator COI:GQ329635 18/10/2006 Female Open montane stream, D2/D3:None riffles, glides, deposition D7: GQ329655 zones 16S: None Syncordulia venator COI:GQ329634 11/11/1999 Female Open montane stream, D2/D3: GQ329639 boulders D7: GQ329654 16S: GQ329646 Syncordulia venator COI:GQ329633 16/11/2006 Male Open montane stream, D2/D3:None boulders D7: None 16S: GQ329645 Syncordulia venator COI:GQ329632 16/11/2006 Male Open montane stream, D2/D3:None boulders D7: GQ329653 16S: GQ329643 Syncordulia venator COI: None 14/11/2006 Female Open montane stream, D2/D3:None boulders D7: None 16S: GQ329644 Syncordulia serendipator COI:GQ329631
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