Biodiversity: time for detailed local studies, R Rakauskas

Tags: biodiversity studies, Lithuania, organisms, samples, Aphis oenotherae, Poland, biodiversity, local studies, distribution areas, Lithuania Biodiversity studies, Ecological diversity, populations, Vilnius University, organismal diversity, Potsdam S. Korea Sandomie Sicily Siedlce Skirgisk Slovakia Swinoujs Toronto Treviso Valakamp Wales Berkeley, aphid species, conservation of biodiversity, nature protection, systematics and biodiversity, systematics, Gliwice, Poland, morphological analysis, Katowice, Poland
Content: BIOLOGIJA. 2006. Nr. 1. P. 91­95
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Biodiversity: time for detailed local studies
R. Rakauskas Department of Zoology, Vilnius University, Ciurlionio 21/27, Vilnius, LT 03001, Lithuania
Biodiversity studies are not only listings of species in local, regional and worldwide faunas and floras. Infraspecific variation might appear an even more exciting field for biodiversity studies. It is common knowledge that species (especially those prospering ones) have large distribution areas. Individuals, demes and populations commonly inhabit distribution areas of their own species unevenly. Remarkable variation of environmental conditions is usually present inside these distribution areas, causing specific trends of adaptive evolution inside the same species, resulting in fixation of locally specific character states, and enforcing designation of various infraspecific units (biotypes, ecotypes, races, subspecies etc.) inside the species. Therefore, detailed ecological, morphological and molecular studies of local populations seem to be urgent for a better understanding of biodiversity and natural management. All this knowledge is summarized by systematics, and might also result in a critical revision of the taxonomic status of a species under analysis. The above statements are illustrated by an example of aphid biodiversity and systematics studies in Lithuania. Key words: biodiversity, organismal diversity, infraspecific structure, Aphis oenotherae, Lithuania
INTRODUCTION The common understanding of biodiversity is rather narrow, including only diversity within species, between species and that of ecosystems. Regionally, biodiversity is usually restricted as conduction of lists of local faunas and floras, with the main attitude to their conservation [1, 2]. The aim of this paper is to draw greater attention and encourage studies in the other fields of biodiversity in Lithuania, with the focus on infraspecific variation and emphasis on the complex studies of local populations. An example will be from the taxonomic studies of aphid species Aphis oenotherae Oestlund, 1887 (Hemiptera, Sternorrhyncha: Aphididae). biological diversity ­ concepts, definitions and common understanding From the numerous definitions of the term Biological diversity, or simply Biodiversity [3], the most widely accepted seems to be the following: "Biological diversity or biodiversity is `the variety of life', and refers collectively to variation at all levels of biological organization" [1]. This variety of life is expressed in a multiplicity of ways and is therefore divided between different key elements (building blocks) in some handbooks. For example, Gaston & Spicer [1] have presented three principle elements of biodiversity. Genetic diversity concerns nucleotides, genes,
chromosomes, individuals and populations. Organismal diversity includes individuals, populations, infraspecific taxa, species, genera, families, phyla, and domains (kingdoms). Ecological diversity starts with populations, and continues with niches, habitats, ecosystems, landscapes, bioregions to biomes. Noticeably, population level is present in all building blocks (elements) of biodiversity. This is inevitable, because a population is an elementary unit of biological evolution; it is the smallest unit of life, being capable to evolve on its own [4]. Once having accepted the above-mentioned concept of biodiversity, one can argue that biology is the science of biodiversity, because biology is a scientific study of living organisms. Naturally, such a broad concern of biodiversity provokes different applications of the term, including the specialised and narrow ones. Article 2 of the Convention on Biological Diversity (CBD) depicts biodiversity as variability among living organisms from all ecosystems and the ecological complexes of which they are part, including diversity within species, between species and of ecosystems [5]. One can argue that such understanding excludes the genetic block of biodiversity. On the other hand, in the frame of the Convention on Biological Diversity (signed under the auspices of United Nations in Rio de Janeiro in 1992), biodiversity is concerned not on its own, but with the objective of "the conservation of biological diversity, the sustainable use of its compo-
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nents..." (Article 1 of CBD). Therefore, in the common knowledge, the term biodiversity is usually associated with the conservation and nature protection. This results in an evil practice when decision makers and budget keepers prefer funding the conservation of biodiversity (e.g., nature protection and sustainable development) rather than the study of biodiversity (e.g., taxonomy). Of course, stocktaking of the local and worldwide species lists, arranging the checklists of faunas and floras is an important and primeval task concerning biodiversity studies, as is the protection of biodiversity. Nonetheless, deeper studies of local populations are of no less importance, especially when concerning the need of the conservation policy. Biodiversity and systematics When dealing with biodiversity, one should remember systematics ­ "the scientific study of the kinds and diversity of organisms and of any and all relationships among them" ([6]: 7). The following passage of Mayr [7] clearly explains the specific role of systematics in biodiversity studies. Systematics is unique among the biological sciences in its dominant concern with diversity. In all subdivisions of functional biology the main concern is with basic processes and mechanisms shared by all or most organisms. Hence the reductionist tendency at the cellular and molecular levels ­ the endeavor to reduce everything to common denominators, i.e. to penetrate to the universal building stones. Biology would become very one-sided if all biologists were to share this objective. It is the student of systematics who helps to restore the balance by his interest in and insistence on uniqueness. One of the major preoccupations of systematics is to determine, by comparison, what the unique properties of every species and higher taxon are. Another is to determine what properties certain taxa have in common with each other, and what the biological causes for the differences or shared characters are. Finally, it concerns itself with variation within taxa. Systematics helps to build proper classifications, and classification makes the organic diversity accessible to the other biological disciplines. Systematics deals with populations, species, and higher taxa. No other branch of biology occupies itself in a similar manner with this level of integration in the organic world. It not only supplies urgently needed information about these levels but, more important, it cultivates a way of thinking, a way of approaching biological problems which is tremendously important for the balance and well-being of biology as a whole. There is a need for someone to stress the diversity of the living world, the most truly biological quality of organisms. However legitimate the study is of that which organisms have in common (much of it being the physics and chemistry of organisms), it is equally legitimate to study the uni-
que characteristics of taxa at all levels down to the subspecies. Therefore, one can hardly discriminate between systematics and biodiversity, because systematics endeavors to order the rich diversity of the Living World (biodiversity) and to develop methods and principles to make this task possible. When supporting systematics, we support biodiversity studies. Naturally, biodiversity studies provide material for systematics and taxonomy. An example from Lithuania ­ aphid biodiversity studies When performing studies of aphid (Hemiptera, Sternorrhyncha: Aphididae) fauna in Lithuania, a new for the local fauna aphid species ­ Aphis oenotherae Oestlund, 1887 ­ has been found in 2002 in the surroundings of Vilnius [8]. Later on, A. oenotherae appeared to be widespread throughout Lithuania and was recorded in Belarus also [9]. The fact would remain of minor importance from the viewpoint of biodiversity studies (just one more country in Europe received one more aphid species already available in other European countries), unless the detailed studies of the Lithuanian clones of A. oenotherae. A. oenotherae has been originally described from Minnesota (USA [10]), later reported as widespread over the other states of USA also [11]. It is supposed as being introduced to Europe at the end of the last century, and has already been reported from Italy [12], Spain [13], United Kingdom [14], Germany [15], southern Poland [16], also Czech Republic and Slovakia (Holman 2000, personal communication). Patch [17] provided an experimental evidence for the Nearctic populations of A. oenotherae (at the time under the name of A. sanborni) being holocyclic species, alternating between Ribes spp. (currants and gooseberries) and Epilobium herbs (Onagraceae, the same plant family that the genus Oenothera belongs to). Therefore, Nearctic populations of A. oenotherae are supposed to be a holocyclic heteroecious species alternating between Ribes spp. and Onagraceous herbs. In Europe, A. oenotherae is known to be anholocyclic (having no bisexual reproduction), inhabiting various species of Oenothera in the field [18, 14]. Mьller [15] has reported on successful transfer experiments to other herbaceous hosts of the plant family Onagraceaea (Godetia, Gaura, Epilobium, Fuchsia, Chamaenerium and Clarkia). Thus, the European populations of A. oenotherae are reported as being anholocyclic ones [18, 14], i. e. they reproduce by parthenogenesis only. In Lithuania, we have found A. oenotherae heavily infesting Oenothera biennis and other species of the genus Oenothera. The aphids cause severe leaf roll and deformation of the stem and inflorescences of their hosts. Up to 80% of Oenothera biennis plants appear to be infested in some places. We failed to find A. oenotherae inhabiting Epilobium, Chamaenerium or other plants of the family Onagraceae in
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Table. Aphis oenotherae samples used for the morphological analysis when constructing the cladogram presented in Fig. 1. Oe = Oenothera
Abbreviated Label data in Fig. 1
Label hostplant
Calif Cantabri Colorado Druskini Gliwice Grodno Hawaii Katowice London Lublin Manitoba Moravia NCarolin Potsdam S. Korea Sandomie Sicily Siedlce Skirgisk Slovakia Swinoujs Toronto Treviso Valakamp Wales
Berkeley, California, USA, 1963.10.30 Comillas, Santander, Cantabria, Spain, 1981.05.24 Denver, Colorado, USA, 1925.04.12 Druskininkai, Lithuania, 2002.08.02, No 8 Przechliebie, Gliwice, Poland, 2002.06.25, No 4 Siabryn'-Ivjevsk, Grodno, Belorus', 2002.08.29, No 02-323 Haleakala, Maui, Hawaii, 1999.04.02, No 7199 Katowice, Poland, 1999.07.19, No Chelsea, London, Great Britain, 1992.07.09, No 6106 Gnojno, Lublin, Poland, 2002.06.27, No 5 Spruce Woods, Manitoba, Canada, 1973.07.26, No V 75-26-7 Bzenec, Moravia, Czech Rep. 1984.07.04, No 18 726-7 Wrightsvile Beach, North Carolina, USA, 1964.04.25, No 262 Kleinmachnow- Potsdam, Germany 1988,06,08 No 20137 Pyomg Chang Kun, Bong Pyomg Myen, South Korea, 1999.06.03, No 99Ho652 Dunkowice, Sandomierz, Poland, 2000.07.21, No 52/20 Zafferana-Catania, Sicily, 1977.09.21 Siedlce, Poland, 2002.06.26, No 2 Skirgiskes, Vilnius region, Lithuania, 2002.10.09, No 02-108 Chotin, Slovakia, 1984.06.25, No 18512+18520 Uznam, Swinoujscie, Poland, 1972,08,15, No 4013 Toronto Ravine, Ontario, Canada, 1968.08.31 Salgareda, Treviso, Italy, 2000.09.04, No 87/20/582 Valakampiai-Vilnius, Lithuania, 2002.07.05, No 02-73 Mid Glamorgam, S. Wales, Great Britain, 1992.07
Oe. sp. Oe biennis Oe. biennis Oe casimiri Oe. subterminalis Oe biennis Oe. sp. Oe. flaemingina Oe. biennis gr.cult Oe. oakesiana Oe. parviflora Oe. biennis Oe. sp. Oe. biennis Oe. odorata Oe biennis Oe. stricta Oe. brevihypanthialis Oe biennis Oe biennis Oe biennis Oe. biennis Oe. stucchii Oe biennis Oe. sp.
Lithuania in the field, despite special searches. In the pot cages, when having no possibility to colonize Oenothera plants, aphids fed on Epilobium adenocaulon for several parthenogenetic generations. Clonal aphids (both winged and apterous ones) refused to accept cultivated black and red currants as host plants. Winged males and apterous oviparous females appeared in the field and in the pot cages on the Oenothera plants from the beginning of October, and the winter eggs were laid on the rosette leaves of first-year Oenothera plants. Thus, A. oenotherae is monoecious holocyclic on Oenothera spp. (probably also on Epilobium spp.) in Lithuania. This is a new knowledge concerning the life cycle of this aphid species in Europe and Nearctics. Morphologically, aphids of the genus Aphis collected on Oenothera biennis in Belarus and Lithuania appeared to be indistinguishable from A. oenotherae samples from other places of the species distribution area: their antennae and siphunculi were short (relatively to body length) and marginal tubercles on the abdominal segments II­VI were absent [9]. The STATSOFT computer programme STATISTICA for WINDOWS (Kernel release 5.5 A) has been used for the cladistic analysis of 25 samples of A. oenotherae from 12 countries (apterous
viviparous females) exploiting fifteen morphological characters. These were the lengths (in mm) of: antennal segments III­V; articular width of antennal segment III; the basal part of antennal segment VI; cauda; longest hair on antennal segment III; processus terminalis on antennal segment VI; siphunculus; second segment of hind tarsus; apical rostral segment; and maximum width of antennal segment III. The counts were of: hairs on cauda; additional hairs on the apical rostral segment; marginal tubercles on abdominal segments II­VI. Data concerning aphid samples are presented in Table. Cladistic analysis based on the above-listed morphological characters did not reveal any morphological specificity of Lithuanian samples (Fig. 1). Lithuanian samples appeared in two main separate clades. Phylogenetic analysis of 34 samples of 8 species of the genus Aphis L. (including 10 samples of A. oenotherae from Lithuania, Poland and South Korea) based on maximum parsimony analysis of the studied region of nuclear sequence of EF 1 showed A. oenotherae to be well separated from the other European species of the subgenus Bursaphis of the genus Aphis (for details, see [19]). Again, Lithuanian samples appeared in separate clades, together with samples from S. Korea and Poland (Fig. 2).
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For the present, Lithuanian populations of A. oenotherae do not seem to be much different in their morphology and sequences of the studied fragments of EF 1 when compared with those from S. Korea and Poland. Nonetheless, the specificity of their life cycle (holocycle and monoecy on Oenothera spp.) suggests them to be different from the Nearctic representatives of A. oenotherae. Although similar morphologically, Nearctic populations of A. oenotherae should be checked in their molecular features. This might result in the description of aphid species new for science. CONCLUSIONS 1. Biodiversity studies are not only the stocktaking of the local and worldwide species and arranging the checklists of faunas and floras. Deeper studies of local populations (infraspecific and genetic diversity) are of equal importance. 2. Biological systematics and biodiversity are inseparable, because systematics endeavor to order the rich diversity of the Living World (biodiversity) and to develop methods and principles to make this task possible. Funding of systematic studies equals the support for biodiversity studies. 3. A complex studies of the Lithuanian populations of A. oenotherae (Hemiptera, Sternorhyncha: Aphididae) revealed them to be clearly distinct in their host specificity and life cycle (monoecious holocyclic on Oenothera spp.) from the European (anholocyclic on Oenothera spp) and Nearctic (heteroecious holocyclic alternating between Ribes spp. and Onagraceae, including Oenothera spp.) populations of the same species.
Linkage Distance
TORONTO MANITOBA SICILY TREVIZO LUBLIN SKIRGISK GLIWICE SLOVAKIA NCAROLIN COLORADO KATOWICE CALIF SIEDLCE CANTABRI WALES MORAVIA SWINOUJS SANDOMIE DRUSKINI S.KOREAI VALAKAMP GRODNO POTSDAM HAWAII LONDON
8 7 6 5 4 3 2 1 0 Fig. 1. Hierarchical tree plot (Euclidean distance, complete linkage) of 25 samples of Aphis oenotherae from 12 countries based on fifteen morphological characters of apterous viviparous females. Arrows indicate samples from Lithuania. Detailed information on aphid samples used is presented in Table LT PL Korea
Fig. 2. Maximum parsimony tree based on the studied region of EF 1a in the genus Aphis L. Bootstrap support based on 1000 replicates is indicated for nodes with greater than 50% support. Samples of A. oenotherae from Lithuania are marked as LT, from Poland as PL, from South Korea as Korea (after [19])
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ACKNOWLEDGEMENTS Author wants to express his sincere thanks to S. Lee (Suwon, Korea) for providing aphid material. Kind assistance of J. Holman, J. Havelka, K. Spitzer, J. Jaros (Ceskй Budйjovice, Czech Republic), W. Wojciechowski, A. Czylok and P. Wegierek (Katowice, Poland) during field sampling cannot be overestimated. Aphid collecting in the Czech Republic has been enabled due to financial support from Czech Academy of Sciences ­ Lithuanian Academy of Sciences Exchange Programme. This work was partially supported by the Norwegian Centre for international cooperation in Higher Education (grant CCP 03/02: ENLINO Master program network) and Lithuanian State Science and Studies Foundation Grant No. C 03056. Received 1 December 2005 Accepted 15 February 2006
10. Oestlund OW. Geological and natural history Survey of Minnesota 1887; 4: 1­99. 11. Palmer MA. Aphids of the Rocky Mountain region. Denver, Thomas Say Foundation, 1952. 12. Barbagallo S, Stroyan HLG. Frust Entom N. S. 1980; 3: 1­182. 13. Nieto Nafria JM, Mier Durante MP. Bolm Soc Port Ent 1985; 3: 115­25. 14. Martin JH. Entomologist's Gazette 2000; 51: 97­105. 15. Muller FP. Entomol Nachr 1974; 18: 129­33. 16. Szelegiewicz H. Ann Zool 1976; 33(13): 217­27. 17. Patch EM. Maine Agr Expt Sta Bull 1927; 336: 1­8. 18. Heie OE. The Aphidoidea (Hemiptera) of Fennoscandia and Denmark. III Leiden-Copenhagen, E. J. Brill / Scandinavian Science Press, 1986. 19. Turcinavicien J, Rakauskas R, Pedersen BV. Eur J Entomol 2006; (in press). R. Rakauskas
References 1. Gaston KJ, Spicer JI. Biodiversity: an Introduction. Oxford, Blackwell Publishing, 2004. 2. Puplesis R. Pasaulio biologin vairov. Kaunas, Lutut, 2002. 3. DeLong DC. Wildlife Society Bull 1996; 24: 738­49. 4. Ridley M. Evolution. Oxford, Blackwell Publishing, 2004. 5. Convention on biological diversity (with annexes). Conc- luded at Rio de Janeiro on 5 June 1992. UN Treaty Series No 30619, 1993. 6. Simpson GG. Principles of animal taxonomy. New York, Columbia University Press, 1961. 7. Mayr E. Principles of systematic zoology. New York, McGraw-Hill, 1969. 8. Rakauskas R. Ekologija (Vilnius) 2004; 1: 1­4. 9. Buga SV, Rakauskas R. Acta Zool Lituanica 2003; 13(4): 396­402.
BIOVAIROV: METAS VYKDYTI ISSAMIUS LOKALIUS TYRIMUS Santrauka Biovairov turi bti suvokiama ne vien tik kaip regioniniai ir viso pasaulio organizm rsi srasai. Vidursin vairov yra ne maziau svarbus biovairovs elementas. Rsys uzima nemazas teritorijas netolygiai jose pasiskirstydamos. vairios aplinkos slygos rsies areale lemia skirtingas (kartais priesingas) tos rsies populiacij evoliucijos kryptis, atsiranda didel vairov rsi viduje: ekotipai, morfotipai, rass, porsiai ir pan.. Issams vietini populiacij tyrimai tiesiog btini siekiant geriau suvokti biologin vairov bei j racionaliai valdyti. Biologin sistematika siame kontekste tampa svarbiu teoriniu ir praktiniu biovairovs tyrimo rankiu. Sie teiginiai straipsnyje iliustruojami pavyzdziais apie amar biovairovs ir sistematikos tyrimus Lietuvoje.

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