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The dispersal–vicariance pendulum and butterfly biogeography
Stockholm University, Faculty of Science, Department of Zoology.
2009 (English)Doctoral thesis, comprehensive summary (Other academic)
Abstract [en]

The relative importance of dispersal and vicariance in speciation is a subject of long standing debate. The earliest historical biogeographers invoked dispersal to explain disjunct distributions. With the advent of phylogenetic systematics and the acceptance of plate-tectonic theory, vicariance gained prominence and dispersalist explanations were increasingly rejected in favour of the former. This led to a new paradigm termed ‘vicariance biogeography’. The quintessence of vicariance biogeography is the consideration of vicariance as the null hypothesis in explanations of disjunct distributions. The notion of vicariance being the predominant force in allopatric speciation started gaining increasing acceptance in the biogeographic community. This also came to be reflected in analytical methods, many of which are biased towards vicariant inferences. However, the recent past has seen this vicariance-dominated view being confronted by a suite of studies demonstrating that dispersal has played a vital role in speciation and is equally important, if not more. In this thesis, I have studied the historical biogeography of nymphalid butterflies (Family Nymphalidae) in the genus Junonia and two subtribes - Coenonymphina and Mycalesina. Junonia is found in all major zoogeographic regions apart from the Palaearctic. Coenonymphina is found in the Holarctic, Neotropical and Australasian regions. Mycalesina is found all over the Old World tropics. The results in the thesis indicate that dispersal has played a crucial role in the diversification of these groups, while there is little evidence for vicariance in any group. I also critique Dispersal-Vicariance analysis, the widely used analytical method in historical biogeography that is based on the principle of parsimony. I use simulated data to highlight various sources of error when using the method, and suggest ways that may help increase the realism of biogeographic inferences.

Place, publisher, year, edition, pages
Stockholm: Department of Zoology, Stockholm University , 2009. , 12 p.
Keyword [en]
Junonia, Coenonympha, Coenonymphina, Mycalesina, historical biogeography, vicariance, dispersal, DIVA
National Category
Natural Sciences
Research subject
Animal Ecology
Identifiers
URN: urn:nbn:se:su:diva-29505ISBN: 978-91-7155-936-4 (print)OAI: oai:DiVA.org:su-29505DiVA: diva2:233837
Public defence
2009-10-09, Lilla hörsalen, Naturhistoriska riksmuseet, Frescativägen 40, Stockholm, 10:00 (English)
Opponent
Supervisors
Note
At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 3: Manuscript. Paper 4: In press. Paper 5: In press.Available from: 2009-09-17 Created: 2009-09-02 Last updated: 2009-09-04Bibliographically approved
List of papers
1. Out-of-Africa origin and dispersal mediated diversification of the butterfly genus Junonia (Nymphalidae: Nymphalinae)
Open this publication in new window or tab >>Out-of-Africa origin and dispersal mediated diversification of the butterfly genus Junonia (Nymphalidae: Nymphalinae)
2007 (English)In: Journal of Evolutionary Biology, ISSN 1010-061X, Vol. 20, no 6, 2181-2191 p.Article in journal (Refereed) Published
Abstract [en]

The relative importance of dispersal and vicariance in the diversification of taxa has been much debated. Within butterflies, a few studies published so far have demonstrated vicariant patterns at the global level. We studied the historical biogeography of the genus Junonia (Nymphalidae: Nymphalinae) at the intercontinental level based on a molecular phylogeny. The genus is distributed over all major biogeographical regions of the world except the Palaearctic. We found dispersal to be the dominant process in the diversification of the genus. The genus originated and started diversifying in Africa about 20 Ma and soon after dispersed into Asia possibly through the Arabian Peninsula. From Asia, there were dispersals into Africa and Australasia, all around 5 Ma. The origin of the New World species is ambiguous; the ancestral may have dispersed from Asia via the Beringian Strait or from Africa over the Atlantic, about 3 Ma. We found no evidence for vicariance at the intercontinental scale. We argue that dispersal is as important as vicariance, if not more, in the global diversification of butterflies.

Keyword
butterfly evolution, DIVA, geo-dispersal, 'Out of Africa', taxon pulses, vicariance
Identifiers
urn:nbn:se:su:diva-19248 (URN)10.1111/j.1420-9101.2007.01425.x (DOI)000250298300011 ()
Available from: 2007-10-30 Created: 2007-10-30 Last updated: 2009-09-04Bibliographically approved
2. Phylogeny and biogeography of Coenonympha butterflies (Nymphalidae: Satyrinae) – patterns of colonization in the Holarctic
Open this publication in new window or tab >>Phylogeny and biogeography of Coenonympha butterflies (Nymphalidae: Satyrinae) – patterns of colonization in the Holarctic
2009 (English)In: Systematic Entomology, ISSN 0307-6970, Vol. 34, no 2, 315-323 p.Article in journal (Refereed) Published
Abstract [en]

We studied the historical biogeography of a group of butterflies in the Holarctic region belonging to the genus Coenonympha (Nymphalidae: Satyrinae: Coenonymphina), based on a phylogenetic hypothesis estimated from three genes. The genus is distributed mainly in the Palaearctic region, with two species extending into the Nearctic region. The tree is generally well supported and shows that Coenonympha is paraphyletic with respect to Lyela (syn.n.) and Triphysa (syn.n.), and we hence synonymize the latter two with Coenonympha. Within Coenonympha we identify three species groups, the tullia, glycerion and hero groups. The North American tullia exemplars are not sister to the Eurasian ones. A diva analysis indicates that the ancestor of the group was present in the Central Palaearctic or Central Palaeartic + Western Palaearctic or Central Palaearctic + Eastern Palaearctic. We conclude that the most likely origin of extant members of Coenonympha was in the Central Asian mountains. The tullia and hero groups started diverging in Europe following dispersal into the region. There have been two independent colonizations into Africa. The drying up of the Mediterranean during the Messinian period probably played an important role, allowing colonization into the Mediterranean islands and Africa.

Identifiers
urn:nbn:se:su:diva-29537 (URN)10.1111/j.1365-3113.2008.00453.x (DOI)000264374300009 ()
Available from: 2009-09-03 Created: 2009-09-03 Last updated: 2009-09-04Bibliographically approved
3. Phylogenetics and biogeography of a spectacular Old World radiation of grass feeding butterflies: the subtribe Mycalesina (Lepidoptera: Nymphalidae: Satyrini)
Open this publication in new window or tab >>Phylogenetics and biogeography of a spectacular Old World radiation of grass feeding butterflies: the subtribe Mycalesina (Lepidoptera: Nymphalidae: Satyrini)
Show others...
2010 (English)In: BMC Evolutionary Biology, ISSN 1471-2148, Vol. 10, 172-185 p.Article in journal (Refereed) Published
Abstract [en]

Background: Butterflies of the subtribe Mycalesina (Nymphalidae: Satyrinae) are important model organisms in ecology and evolution. This group has radiated spectacularly in the Old World tropics and presents an exciting opportunity to better understand processes of invertebrate rapid radiations. However, the generic-level taxonomy of the subtribe has been in a constant state of flux, and relationships among genera are unknown. There are six currently recognized genera in the group. Mycalesis, Lohora and Nirvanopsis are found in the Oriental region, the first of which is the most speciose genus among mycalesines, and extends into the Australasian region. Hallelesis and Bicyclus are found in mainland Africa, while Heteropsis is primarily Madagascan, with a few species in Africa. We infer the phylogeny of the group with data from three genes (total of 3139 bp) and use these data to reconstruct events in the biogeographic history of the group.

Results: The results indicate that the group Mycalesina radiated rapidly around the Oligocene-Miocene boundary. Basal relationships are unresolved, but we recover six well-supported clades. Some species of Mycalesis are nested within a primarily Madagascan clade of Heteropsis, while Nirvanopsis is nested within Lohora. The phylogeny suggests that the group had its origin either in Asia or Africa, and diversified through dispersals between the two regions, during the late Oligocene and early Miocene. The current dataset tentatively suggests that the Madagascan fauna comprises two independent radiations. The Australasian radiation shares a common ancestor derived from Asia. We discuss factors that are likely to have played a key role in the diversification of the group.

Conclusions: We propose a significantly revised classification scheme for Mycalesina. We conclude that the group originated and radiated from an ancestor that was found either in Asia or Africa, with dispersals between the two regions and to Australasia. Our phylogeny paves the way for further comparative studies on this group that will help us understand the processes underlying diversification in rapid radiations of invertebrates.

National Category
Biological Systematics
Research subject
Animal Ecology
Identifiers
urn:nbn:se:su:diva-29539 (URN)10.1186/1471-2148-10-172 (DOI)000279831400001 ()
Funder
Swedish Research Council
Note
authorCount :6Available from: 2009-09-03 Created: 2009-09-03 Last updated: 2010-12-14Bibliographically approved
4. Phylogenetics of Coenonymphina (Nymphalidae Satyrinae) and the problem of rooting rapid radiations
Open this publication in new window or tab >>Phylogenetics of Coenonymphina (Nymphalidae Satyrinae) and the problem of rooting rapid radiations
Show others...
2010 (English)In: Molecular Phylogenetics and Evolution, ISSN 1055-7903, E-ISSN 1095-9513, Vol. 54, no 2, 386-394 p.Article in journal (Refereed) Published
Abstract [en]

We report a rapid radiation of a group of butterflies within the family Nymphalidae and examine some aspects of popular analytical methods in dealing with rapid radiations. We attempted to infer the phylogeny of butterflies belonging to the subtribe Coenonymphina sensu lato using five genes (4398bp) with Maximum Parsimony, Maximum Likelihood and Bayesian analyses. Initial analyses suggested that the group has undergone rapid speciation within Australasia. We further analyzed the dataset with different outgroup combinations the choice of which had a profound effect on relationships within the ingroup. Modelling methods recovered Coenonymphina as a monophyletic group to the exclusion of Zipaetis and Orsotriaena, irrespective of outgroup combination. Maximum Parsimony occasionally returned a polyphyletic Coenonymphina, with Argyronympha grouping with outgroups, but this was strongly dependent on the outgroups used. We analyzed the ingroup without any outgroups and found that the relationships inferred among taxa were different from those inferred when either of the outgroup combinations was used, and this was true for all methods. We also tested whether a hard polytomy is a better hypothesis to explain our dataset, but could not find conclusive evidence. We therefore conclude that the major lineages within Coenonymphina form a near-hard polytomy with regard to each other. The study highlights the importance of testing different outgroups rather than using results from a single outgroup combination of a few taxa, particularly in difficult cases where basal nodes appear to receive low support. We provide a revised classification of Coenonymphina; Zipaetis and Orsotriaena are transferred to the tribe Eritina.

National Category
Natural Sciences
Identifiers
urn:nbn:se:su:diva-29524 (URN)10.1016/j.ympev.2009.08.012 (DOI)000273758200007 ()19686856 (PubMedID)
Available from: 2009-09-03 Created: 2009-09-03 Last updated: 2014-10-28Bibliographically approved
5. Use of dispersal–vicariance analysis in biogeography – a critique
Open this publication in new window or tab >>Use of dispersal–vicariance analysis in biogeography – a critique
2010 (English)In: Journal of Biogeography, ISSN 0305-0270, E-ISSN 1365-2699, Vol. 37, 3-11 p.Article in journal (Refereed) Published
Abstract [en]

Aim Analytical methods are commonly used to identify historical processes of vicariance and dispersal in the evolution of taxa. Currently, dispersal-vicariance analysis implemented in the software diva is the most widely used method. Despite some recognized shortcomings of the method, it has been treated as error-free in many cases and used extensively as the sole method to reconstruct histories of taxa. In light of this, an evaluation of the limitations of the method is needed, especially in relation to several newer alternatives. Methods In an approach similar to simulation studies in phylogenetics, I use hypothetical taxa evolving in specific geological scenarios and test how well diva reconstructs their histories. Results diva reconstructs histories accurately when evolution has been simple; that is, where speciation is driven mainly by vicariance. Ancestral areas are wrongly identified under several conditions, including complex patterns of dispersals and within-area speciation events. Several potentially serious drawbacks in using diva for inferences in biogeography are discussed. These include the inability to distinguish between contiguous range expansions and across-barrier dispersals, a low probability of invoking extinctions, incorrect constraints set on the maximum number of areas by the user, and analysing the ingroup taxa without sister groups. Main conclusions Most problems with inferences based on diva are linked to the inflexibility and simplicity of the assumptions used in the method. These are frequently invalid, resulting in spurious reconstructions. I argue that it might be dangerous to rely solely on diva optimization to infer the history of a group. I also argue that diva is not ideally suited to distinguishing between dispersal and vicariance because it cannot a priori take into account the age of divergences relative to the timing of barrier formation. I suggest that other alternative methods can be used to corroborate the findings in diva, increasing the robustness of biogeographic hypotheses. I compare some important alternatives and conclude that model-based approaches are promising.

 

Identifiers
urn:nbn:se:su:diva-29536 (URN)
Available from: 2009-09-03 Created: 2009-09-03 Last updated: 2011-01-10Bibliographically approved

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