Phylogenetic Systematics and Historical Biogeography of Malesian
Calicnemiine Damselflies (Odonata, Platycnemididae)
Gassmann, Dirk
Citation
Gassmann, D. (2005, October 19). Phylogenetic Systematics and Historical Biogeography
of Malesian Calicnemiine Damselflies (Odonata, Platycnemididae). Retrieved from
https://hdl.handle.net/1887/9758
Version:
Corrected Publisher’s Version
License:
Licence agreement concerning inclusion of doctoral thesis in the
Institutional Repository of the University of Leiden
INTRODUCTION General
Based upon the late recognition of both Alfred Wegener's theory of plate tectonics (1915) and Willi Hennig's phylogenetic systematics (1950, 1966), as well as by the influence of Leon Croizat's panbiogeography (1958) and the recent developments in analytical and computerized biogeographal methodology, the last decades led to a considerable increase of historical biogeographical studies on various groups of organisms, using different types of characters. The melange between a steadily growing geological-tectonical knowledge of the earth's surface on one hand, and the advancement of methodology in comparative historical biogeography on the other hand, has proven to be fruitful.
Certainly, the recognition that taxa and areas may evolve together, and its implementation in biogeographical research, was one of the foremost achievements in historical biogeographic research in the second half of the last century. While before Croizat (e.g.: 1958) disjunct distributions of presumably related taxa were predominantly considered the result of long distance dispersal, this view slowly shifted towards the vicariance approach.
The proponents of 'vicariance biogeography' were not always free of dogmatism, when rejecting dispersal hypotheses as 'old-fashioned'. Also, the 'old' (non-analytical) biogeography was occasionally dismissed as 'narrative', while it could certainly better be called 'inductive' instead. On the contrary, the limits of area-cladistic biogeography have been shown by several authors (e.g. Cracraft, 1988; Gittenberger, 1990; Polhemus, 1996), as for instance the difficulties to cope with complicated reticulate patterns, with multiple histories, or the potential inconstancy of its operational units, the areas. It appears that, after a phase of overestimation of vicariance as a factor in radiation and in speciation within monophyletic groups, the importance of long-distance-dispersal has been reappraised recently (Cook & Crisp, 2005; De Queiroz, 2005).
The last decade witnessed an increased cooperation between biogeographers and geologists focussing on the Southeast Asian region resulting in a series of scientific meetings and publications. Obviously, on the one hand geologists can help biogeographers to formulate, or test, hypotheses on ancestral biotas in the region. On the other hand, the findings of biogeographers can help geologists to falsify or corroborate their ideas on the distribution of former land masses and island arcs. Biologists can now compare their findings with comprehensive geological scenarios for Southeast Asia and the Indo-Pacific provided for the Mesozoic (e.g. Metcalfe, 1996) and especially the Cenozoic (e.g. Hall, 2002).
In the meantime, the phylogeny of various groups of Malesian organisms has been studied by biologists using cladistic methods, thus creating the basis for both revealing individual distribution histories of certain groups as well as seeking general patterns caused by vicariance (e.g. Van Welzen et al., 2003). The latter are mainly determined by geological processes.
Pathways to New Guinea
The damselflies of the subfamily Calicnemiinae are an example of a group of organisms occurring on both sides of Wallace's line, i.e. the border along which Oriental and Australian faunal and floral elements are considered to be in balance. The rich diversity of the Calicnemiinae in the Philippines suggests that the eastern representatives of the group could have reached New Guinea by pathways different from the southern (Gondwanan) route. The present project formed part of a research programme of the Netherlands Science Foundation, including the plant family Euphorbiacae with the genera Baccaurea Lour., Distichirhops Haegens and Nothobaccaurea Haegens (Haegens, 2000) and the cicada subtribes Dundubiina and Cosmopsaltriina (Beuk, 2002). Also the present project aims at revealing some of these 'Pathways to New Guinea' based on the distribution and the phylogeny of calicnemiine damselflies.
The Calicnemiinae
Diversity and distribution
Next to the nominal subfamily Platycnemidinae Yakobson & Bianki, 1905, the Calicnemiinae Fraser, 1957, is the second subfamily of the feather-legged damselflies, family Platycnemididae (Odonata). The quite heterogeneous group is confined to the Old World tropics (Fig. 1). The Calicnemiinae currently comprise about 165 species (Schorr et al., 2004), divided into 22 genera. The distribution range of the subfamily includes tropical and subtropical (southern) Africa, the Madagascan region including the Seychelles, the Arabian Peninsula, northern India and the southern flanks of the Himalayas, Southeast Asia up to the Greater Sunda Islands, the Philippines, and the Papuan region from western New Guinea to the Solomons. The Southeast-Asian and Indo-Pacific regions by far house the highest calicnemiine diversity. The genus Risiocnemis Cowley, 1934, (Philippines, excluding the Palawan region) alone comprises 36 species (Gassmann & Hämäläinen, 2002). With 33 species, 19 of which belonging to the large New Guinean genus Idiocnemis Selys, 1878, (Gassmann 1999, 2000), the Papuan region does not only exhibit a similar diversity, but it also accounts for the broadest range of regional morphological diversity among Calicnemiinae, which triggered the introduction of several monotypic genera from that region.
the distribution area. For details on taxonomy and distribution of the Calicnemiinae, see Gassmann (2005) and Van Tol & Gassmann (in press), i.e. chapters 4 and 5 of this thesis.
Fig. 1: World distribution of Calicnemiinae (smaller island groups not indicated).
Fossil record
According to Nel & Papazian (1990), the family Platycnemididae is only very rarely represented in the fossil record. Apart from that, the identity of the fossil specimens described up to now can be doubted. For instance, the taxonomic status of the Late Oligocene adult specimen from France, that was figured by Nel & Papazian (1990) and placed within Platycnemididae, is far from certain. Only the anterior distal part of a single wing of that specimen is preserved, with only the R3 and IR2 veins complete. General shape and the pterostigma appear 'platycnemidid-like'; however, since the taxonomically important IR3, cubital and anal veins are lacking, it seems not possible to detect any character potentially assignable to the ground pattern of Platycnemididae in the present wing fragment.
author of the present thesis is inclined to place those specimens from amber to the Coenagrionidae.
A few further fossil 'platycnemidid' specimens are mentioned in the literature (Nel & Papazian, 1990; Pfau, 1975); they are scattered throughout different collections and should be reexamined.
Larval characters
The present thesis deals with a phylogenetic analysis of morphological characters of the imagines. However, a study of larval characters would be equally interesting, as in some cases useful characters have already been found to distinguish between genera and apparently even between species (Lieftinck, 1958). Unfortunately, very little information is available on larval characters; larvae of two species at most have been described only from Allocnemis Selys, 1863 (Barnard, 1937), Calicnemia Strand, 1928 (Lieftinck, 1984), Coeliccia Kirby, 1890 (Lieftinck, 1958), Idiocnemis Selys, 1878 (Lieftinck, 1958), and R. (Risiocnemis) Cowley, 1934 (Needham & Gyger, 1939). Larvae of several species of Copera Kirby, 1890 and Platycnemis Burmeister, 1839 (i.e. the subfamily Platycnemidinae) are well known (Lieftinck, 1958). The specific identification of most of the described larvae is suppositional (Lieftinck, 1958). When more extensively studied, larval morphology may also support the monophyly of the family Platycnemididae: in the larval forms known and illustrated, the apices of the caudal lamellae are distinctly pointed instead of rounded or only slightly pointed as in Coenagrionidae.
Preliminary molecular work
Fig.2: Strict consensus of two cladograms derived from parsimony analysis of mitochondrial DNA
sequences (Cytochrome Oxidase II). The two most parsimonious trees differ in the position of the two subgenera of Risiocnemis Cowley: Risiocnemis appears either mono-, or polyphyletic. Representatives of coenagrionid genera are used as outgroup. Rooted this way, Copera and Allocnemis appear terminal in the cladogram. The inclusion of additional taxa and characters is required. [With thanks to S. Jordan for providing the sequences]. Abbreviations: AFR = Africa; FIJI = Fiji Islands; HAW = Hawaii Islands; NG = New Guinea; PHIL = Philippines; SEA = Southeast Asia.
Habitat
The fact that calicnemiine damselflies inhabit dense rainforests, occur in comparatively stable habitats, and have a quite low power of dispersal, makes them a suitable group for historical-biogeographic studies. Accordingly, the degree of endemism in the subfamily is high, both at species- and genus-level (see Chapters 4 and 5).
In contrast, most species of the New Guinean Idiocnemis are found at lower altitudes; those species all belong to the I. inornata species-group (Gassmann, 1999), which even includes one representative recorded from the southern New Guinean lowlands. The representatives of the I. bidentata - group (Gassmann, 2000) can reach an altitude of up to 1800 m. Idiocnemis not only exemplifies intra-generic ecological differences; there is also intra-specific variation in habitat choice. Idiocnemis obliterata Lieftinck, for instance, occurs from 50 m to 1000 m and exhibits altitude-dependant phenotypes: individuals from higher altitudes are larger and darker than specimens from lower altitudes. Idiocnemis inaequidens Lieftinck occurs in Papua New Guinea in comparatively open, stony brooks (Fig. 3a) as well as in smaller, more cryptic, forest streamlets with clayey underground (Fig. 3b, own obs.). Contrarily, I. kimminsi Lieftinck and I. adelbertensis Gassmann appear to prefer very shady creeks where they are inconspicuous due to their cryptic red-brown coloration (own obs.). Lieftinck (1958) describes such a habitat also for I. inornata Selys; thus it seems reasonable to assume that these ecological preferences have developed by the ancestor of the I. inornata species-group.
Fig. 3 a/b: Habitats of Idiocnemis inaequidens in Papua New Guinea: Morobe Province, near Lae (a),
Huon Peninsula, nr. Sattelberg (b). Photos: D. Gassmann.
ACKNOWLEDGEMENTS
Klaas-Douwe B. Dijkstra provided up-to-date information on the distribution of African Platycnemididae.
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