Manual The Rhopalocera of Java: Hesperidae

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Only a small number of works devoted to the various causes and factors of range conjugacy have been published since V. Heptner first defined conjugated ranges. In connection with this, I have considered some new aspects of arealogical interactions of butterflies in the tropical ecosystems of Vietnam and other regions of Southeast Asia Monastyrskii, In particular, the species ranges forming the so-called secondary contact or secondary intergradation zones were referred to as interactive ranges.

In addition, of special interest were connections between the ranges of species forming complexes of inedible models and their mimics. In my opinion, such ranges, referred to as mimetic, represent a class of conjugated ranges. Below, I will consider examples of vicariant, interactive, and mimetic ranges of butterflies distributed in Vietnam, Indochina, and other regions of Southeast Asia with the purpose of showing their possible use in working out scenarios of faunogenesis.


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Vicariant Ranges Speaking of geographic vicariance we mean the nature of conjugacy of the range boundaries of several related taxa and the way in which taxa are replaced depending on natural zonality. When considered separately, the range of each of the vicariant taxa may be classified into one of the above arealogical types and configuration variants: continuous, mosaic, and disjunct ranges. However, the phenomenon of vicariance is already another level of geographic organization of animals associated not so much with the vagility of taxa as with their ecological valency, i.

Butterflies (Lepidoptera, Rhopalocera) - Southern Highlands Nature Map

Thus, vicariance may be defined as replacement of the range of one taxon or a group of taxa of a certain rank by ranges of other taxa of the same rank in similar biotopes of different geographic regions. Replacement of ranges of some species by ranges of other species in the meridional direction in the territory of Vietnam was observed in different taxa and considered in detail in earlier publications Monastyrskii, , ; Monastyrskii and Holloway, In the case of uniform vicariance of Nymphalidae, demonstrated in the autochthonous genera of the subfamily Amathusiinae Aemona Hewitson, Stichophthalma C.

Felder et R. In addition, the range boundaries of the replacing species often coincide with the boundaries of the zoogeographic entities distinguished by me in Vietnam Monastyrskii, , Patterns of range replacement in a number of allochthonous butterfly groups of Indochina are associated with centers of their species diversity. Their diversity centers lie in China, and only occasional species with very small local ranges reach the central and southern mountain regions of the Indochinese Peninsula Fig. On the contrary, ranges of most representatives of the large Indo-Australian genus Arhopala Boisduval Lycaenidae, Theclinae are concentrated in the southern regions of Vietnam, whereas the number of replacing species declines sharply on going north.

Vicariant ranges provide an important methodological tool since they reveal the directions of evolutionary processes, i.

Examples of this type of conjugacy can be also effectively used in developing the schemes of biogeographic ranging, which was demonstrated by the example of Vietnam zonality Monastyrskii, , Interactive Ranges The interactive ranges Monastyrskii, are ranges of such species that happened to be separated under the influence of climatic or other factors and then secondarily united already after the development of some specific morphological traits in the isolated populations. Analysis of the butterfly fauna of Southeast Asia, including Vietnam, shows the presence of a considerable number of phenotypically similar species in the tropical ecosystems.

I found over 70 examples of close phenotypic similarity among representatives of the families Papilionidae, Pieridae, and Nymphalidae in the territory of Vietnam. The reasons for such numerous examples of sympatry remain a matter of argument. Distribution of the representatives of the genus complex Zephyrus after Koiwaya, Intensive processes of divergence, i. Fragmentation of the populations of the Indochinese butterfly fauna took place in different geological periods, which is partly confirmed by the varying size of species ranges.

Such species-specific characters as valency and vagility changed under the influence of different factors. Under the conditions of global warming for instance, as a consequence of the latest marine transgression the ranges expanded and their isolated fragments merged forming zones of secondary contact. I do not consider here such a subtle case as the possible hybridization of species after the merging of their ranges Gritsenko et al.

Instead, I am mostly interested in the very fact of the existence of two and more sympatric species, the scale and nature of over- lap of their ranges, and also the possibility of using arealogical interactions in developing the probable faunogenetic scenarios. Such a large and heterogeneous group as butterflies provides examples of different results of the interactions of ranges of the taxa of common origin. In some cases, the ranges of formerly separated related taxa have become close but have not yet acquired an overlap zone.

In other cases, overlapping of ranges and formation of secondary contact zones of different size took place. Interactive ranges of the butterflies of Vietnam; a Elymnias casiphone 1 and E. Ypthima similis Elwes et Edwards and Y. These examples are important since they demonstrate the directions of separation and subsequent convergence of the populations which once belonged to one species. All the pairs listed in the second group may be found simultaneously in the areas of range overlap.

Sometimes the range of one of these species may be considerably reduced and totally included into a wide range of another species; this is observed in such pairs as Elymnias casiphone and E. In other cases, similar species have mosaic ranges, for instance, the mountain species Parantica melaneus and P. The overlap zone is often elongated from north to south but sometimes it has vague and inexplicable outlines. For example, the wide range of Euploea algea Godart Danainae overlaps the narrower range of the sympatric species Eu.

In other cases, the overlap zone is so extended that it is very difficult to understand how and in which direction the ranges of these species changed. A good example is overlapping of ranges of highly phenotypically similar butterflies Pantoporia hordonia Stoll and P.

It may be supposed that the range of the initial species gradually contracted in the course of global cooling and intensive speciation took place in the places of such contractions. Interactive ranges of the butterflies of Vietnam; a Euploea algea 1 and Eu. On the other hand, this configuration may also mean that initial separation took place outside Indochina, yet after the formation of the secondary contact zone and under the influence of the changing climate, for instance, the Holocene warming, both ranges began to expand and reached Indochina in this way, which is manifest in their modern sympatry.

Of special interest are those cases when separation of ranges took place in the territory of Indochina Vietnam , where now both range overlap zones and examples of stable disjunctions can be observed. Analysis of phenotypically similar butterfly species recorded in Indochina showed that many of them originated from other parts of the Indo-Malayan Region Holloway, , and penetrated into the continent and the Indochinese Peninsula long before the cold periods, during which most of them were forced back to the south.

Yet some stenotopic species remained in probable refugia, such as relatively warm areas within the maritime lowlands, or mountain valleys of central and southern Indochina. These isolated areas, like islands, became the main sites of active speciation, which is confirmed by a high concentration of endemics in them Monastyrskii and Holloway, At the same time, the migrating populations managed to preserve their original aspect.

In the course of subsequent warming events this mobile part of the species headed for their initial habitats, creating zones of contact with populations of the refugia which by that time had already become distinct species. This accounts for the numerous examples of sympatry: Euploea orontobates Fruhstorfer and Eu. It is also evident that not all the representatives of the Malayan faunistic element had an opportunity to return; the remainder of their former presence are to be seen in the group of more ancient Malayan endemics: Zeuxidia sapphirus Monastyrskii et Devyatkin, Z.

Examples of pairs of similar related species also occur among the butterfly groups which are of ChineseHimalayan origin; however, they are distinguished by the fact that their ranges practically never overlap.

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The Chinese-Himalayan butterfly species forced southwards by cooling did not return to the northern latitudes after the onset of warming but moved to greater altitudes in the mountain regions of central and southern Indochina where intensive speciation took place. Such a scheme of events explains the appearance and concentration of endemic species and especially subspecies with Chinese-Himalayan connections in these territories. This is an example of the general hypothetical mechanism of species migration during climate fluctuations. The pattern of the prevalent expansion vectors of recent forms also agrees with it.

For example, there are more examples of migration of representatives of the forest flora and fauna to the Arctic Chernov, , , than those of migration of the arctic species to the taiga. If this direction of migrations be taken as the main trend, introduction of the Malayan butterfly species to the continent and their further expansion to the higher latitudes during warming becomes clear.

At the same time, cooling had little effect on retreat of the Malayan species towards the equator since they were preserved in the continental refugia.

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Each consecutive cooling cut off the Malayan species which had penetrated to the continent from the region of autochtonous processes, whereas each warming was accompanied by migration of the next group of Malayan species to the more northern latitudes; this scenario may explain the abundance of similar phenotypes. A good example may be representatives of the large Malayan genus Euploea Fabricius, which could reach Northeast India during the interglacial stages of the Pleistocene.

The regular traits in the migrations change in the range boundaries , separation of taxa, their divergence and subsequent merging of ranges show that these processes are essentially closer to ecological adaptations, i. Mimetic Ranges The mimetic ranges are coinciding ranges of two or more species which are in mimetic relations model— mimic or model—model.

Earlier works on mimicry covered various aspects of this phenomenon: the origin, mechanism, and effectiveness of mimicry, the selective advantage of mimetic polymorphism, and many others Berg, ; Ackery and Vane-Wright, ; Turner, The biogeographic aspects of mimetic coadaptations, in particular to what extent the range boundaries of models and their mimics coincide and how the representatives of both groups are distributed, were much less frequently considered.

Practically all the classical examples of the model— mimic associations can be found among the IndoMalayan butterflies.

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Australian Museum

Main Audio. Hasora Moore, Published in: Watson, E. Parata Moore, synonym. Published in: Moore, F. Hasora Moore, accepted.


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