UNIVERSITY OF PANNONIA GEORGIKON FACULTY FESTETICS DOCRORAL SCHOOL Head of School: Prof. Dr. Angéla Anda
Integrative taxonomic revision of Niphargus spp. and other rare and endemic troglobiont macroinvertebrates from the caves of the Western Mecsek Mts. (South Hungary)
Doctoral (PhD) Theses
DOROTTYA ANGYAL Hungarian Natural History Museum Supervisor: Dr. Előd Kondorosy
KESZTHELY 2015
1. INTRODUCTION, AIMS
Hungarian biospeleological research possessed a rather flourishing epoch during the first half of the last century. Illustrious zoologists of the era, among others Endre Dudich, Elemér Bokor, Antal Gebhardt and Lajos Méhely had started their extensive faunistic, taxonomic and ecological surveys in some of the caves of the Aggtelek Karst and the Mecsek Mts. (e.g. Bokor 1921, Méhely 1924, Dudich 1932, Gebhardt 1934). Between the 1950s and 1970s, still related to the invertebrate zoological survey of the Baradla Cave, mainly taxonomic studies had been published (e.g. Andrássy 1959, Dózsa-Farkas & Loksa 1970, Zicsi 1974). Then, over the past decades the amount of invertebrate zoological research in the Hungarian caves has declined significantly. Bajomi (1977) referred only 49 caves that had been studied in zoological aspect until the end of the 1970s, and this number had only slightly increased until 2010. Due to its protected geographical situation and its isolation, the fauna of the caves situated in the Mecsek Mts. is rather diverse. Numerous endemic species are known from the area. The hypogean habitats of the mountain range with their constant subterranean environment may have played a refugial role during the alternating warmer and colder eras, preserving old lineages of various invertebrate taxa (Angyal et al. 2015). As a result of the intensive zoological survey of the 1920s and 1930s, from the Abaligeti Cave and the Mánfai-kőlyuk Cave – both situated in the Western-Mecsek Mts. – 194 and 159 animal species, respectively had been revealed (Gebhardt 1967). However, during the last decades the character of the two caves had partly changed: the Abaligeti Cave had been developed for the public, while the Mánfai-kőlyuk Cave had been utilized by waterworks. The two caves are known as type localities of several rare and endemic macroinvertebrate taxa too (e.g. Méhely 1924, Soós 1927). Due to the incomplete descriptions and to the specialists’ controversial opinions, taxonomic state of these species and subspecies was rather uncertain. During my research, I was going to resolve these uncertainties using integrative approach in my revisions, based on the combination of traditional methods with modern morphological and molecular taxonomic analyses. As objects of my PhD thesis, seven rare and endemic macroinvertebrate species and subspecies were chosen from the caves of the Western-Mecsek Mts.: the amphipod Niphargus molnari Méhely, 1927 and Niphargus gebhardti Schellenberg, 1934, the aquatic isopod
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Protelsonia hungarica hungarica Méhely, 1924 and Protelsonia hungarica robusta Méhely, 1927, the hydrobiid Bythiospeum hungaricum (Soós, 1927) and Bythiospeum cf. gebhardti (H. Wagner, 1931) and the polydesmid Brachydesmus troglobius Daday, 1889.
In the light of the above, my aims were as follows: to revise the seven above mentioned rare and endemic troglobiont macroinvertebrate species and subspecies and to clarify their taxonomic positions; to contribute to the knowledge on morphology and molecular genetics of the focal taxa with the help of newly applied methods, like scanning electron microscopy, DNA barcoding or phylogenetic analysis; to record the new distributional data of the focal species and subspecies and to make suggestions on their conservation planning, based on evaluation of their rarity and the present condition of their habitats; and to create faunistic lists based on the newly collected aquatic and terrestrial macroinvertebrate material other than the revised species and subspecies.
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2. APPLIED METHODS 2.1 Sampling For my investigations I chose 14 caves of the Western-Mecsek Mts. The previously studied Abaligeti Cave and the Mánfai-kőlyuk Cave were also included. The invertebrate fauna of the further 12 inflow cave, outflow cave and pothole was unknown until the beginning of my research. Between September 2010 and December 2013 I regularly visited the caves for sampling and to make observations. The most frequently used collecting method was ’singling’, which was completed with the use of pitfall tarps in case of the terrestrial fauna, and with bottle traps and leaf litter traps in case of the aquatic fauna. The epicarstic water was collected too, then it was filtered by plankton net. The collected specimens were stored in 70 and 96% ethanol. Identification of the collected material happened by group specialists – in most cases taxonomists of the Hungarian Natural History Museum (HNHM), Department of Zoology – and by me. Specimens were placed in the collections of the HNHM. During the revisions, old museum samples of Stenasellidae and Brachydesmus preserved in the Crustacea and Myriapoda collections of the HNHM were also used. 2.2 Methods of analyses 2.2.1 Morphological studies Niphargus specimens were cooked in 10% KOH solution, rinsed with HCl and washed in distilled water. Cleared exoskeletons were stained with chlorazol black in glycerol, and then dissected under a Leica MZ75 stereomicroscope. Dissected appendages, mouth parts and other organs were mounted in glycerol gelatin. The slides were examined using a Leica DM 1000 light microscope. Drawings were made using a drawing tube attached to the light microscope. Morphological analysis had happened by recording 230 characters on each specimen. Character matrices were compiled and were used for further analysis. Measurements were made using the AnalySIS program package, the computer was connected with a Zeiss Axioscope II light microscope. During the Protelsonia studies same procedure was applied for the dissection and for making of drawings as detailed above. For the morphometric analysis of Bythiospeum specimens, three independent photos were made of each animals.
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Measurement data were recorded by Image J scientific image processing program. Principal Component Analysis and General Linear Model Analysis were performed using the SPSS Statistics software package. Brachydesmus diplopods were studied under a Leica M125 stereomicroscope. Males’ gonopods were dissected and examined at higher magnification. SEM images were made with a HITACHI S-2600 N scanning electron microscope. Dry samples were stuck onto holders and were sputter-coated by gold-palladium. Multilayer photos were shot with Nikon D5200 camera using Mitutoyo M Plan Apo 5X microscope lens.
2.2.2 Molecular taxonomic studies During the molecular studies I performed the laboratory work in the Molecular Taxonomic Laboratory of the HNHM, the Subterranean Biological Laboratory of the University of Ljubljana and in the Molecular Systematic Laboratory of the Natural History Museum Vienna. QIAamp DNA Microcit® and Sigma Aldrich GenElute Mammalian Genomic DNA Miniprep Kit® were used for the DNA extraction. During the Niphargus studies, both mitochondrial (COI) and nuclear (28S rDNA, histone H3) markers were used, while in case of the other taxa only mitochondrial (COI, 16S rRNA) markers were studied. Gene fragments were amplified by polimerase chain reactions. PCR products were checked by gel electrophoresis. Fragments were sequenced in the laboratory of the HNHM and by Macrogen Europe and LGC Genomics. Sequences were edited by BioEdit and were fitted by ClustalW and MAFT 7 programs. Phylogenetic trees and distance matrices were created using MrBayes v3.2 and MEGA 6 software. COI gene fragments of the revised taxa as barcodes were uploaded on the database of National Center for Biotechnology Information.
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3. NEW SCIENTIFIC RESULTS (THESES)
1. Preparing of richly illustrated redescriptions of the amphipod species Niphargus molnari Méhely, 1927 and Niphargus gebhardti Schellenberg, 1934, which agree with the modern Niphargus-taxonomic requirements, and to make their phylogenetic analysis. After the analysis of 230 morphological characters on each specimen, 49 consistently differing characters were found between the two species, which were illustrated by numerous SEM images and drawings. As part of the redescriptions, we compared the main morphological traits of the two species from the Mecsek Mts. with the phylogenetically and geographically close species. After the phylogenetic analysis of 104 European amphipod taxa, we revealed that N. gebhardti belongs to the clade of Central to Eastern European species. Its in sister relationship with N. bihorensis. Other closely related species include Pontoniphargus racovitzai from the Romanian Movile Cave – known for its chemoautotrophic ecosystem – and epikarst-dwelling Southern Slovenian species. We proved that the two species from the Mecsek Mts. are not phylogenetically closely related and they probably colonized the area independently. By comparison of N. gebhardti COI gene fragments from six different caves, we detected that the specimens belong to the same haplotype, which may refer to the existence of genflow through the epikarstic fissures. 2. Clarification of the asellid subspecies Protelsonia hungarica hungarica Méhely, 1924 and Protelsonia hungarica robusta Méhely 1927 by the combination of traditional and modern morphological methods, using old museum samples (HNHM) and newly collected material. Analysis of the collected material revealed two morphologically distinct groups based on consistently differing characters. Differential characters were illustrated with scanning electron micrographs and multilayer photos. I stated that on morphological base, handling P. hungarica hungarica and P. hungarica robusta as two separate subspecies is relevant. Approximately 30% of the specimens collected from the Mánfai-kőlyuk Cave belonged to the ’Abaliget morphotype’. Three possible explanations for this phenomenon are given in the thesis.
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3. Clarification of the degree of relationship between the hydrobiids Bythiospeum hungaricum (Soós, 1927) and Bythiospeum cf. gebhardti (H. Wagner, 1931) by comparison of COI and 16S sequences and by shell morphometric analyses; phylogenetic studies including further species from the Rissooidea superfamily. After the analysis of COI and 16S sequences of hydrobiids from the Abaligeti Cave and the Mánfai-kőlyuk Cave, we revealed 7.05% difference, which – in an other study – meant distinct species in case of a closely related genus. The examined specimens belonged to two haplotypes. Phylogenetic studies revealed that the two species from the Mecsek Mts. are not closely related to the other – mainly Alpine – Bythiospeum species with known sequences. COI distance matrix analysis within the Rissooidea superfamily also showed that the two focal species significantly (in 18-23%) separate from species of other rissooid genera. Morphological distinctness detected by shell morphometric methods supported the molecular genetic results. Similarly to Protelsonia, circa 30% of the specimens collected in the Mánfai-kőlyuk Cave belonged to the ’Abaliget haplotype’, which showed by shell morphometric results too. Four possible explanations for this phenomenon are given in the thesis. 4. Completion of richly illustrated redescription of the polydesmid Brachydesmus troglobius Daday, 1889 based on old museum samples (HNHM) and newly collected material; performing phylogenetic analysis involving further polydesmid species. Fine scale characters were possible to be illustrated in details by scanning electron micrographs and by multilayer photos for the species redescription. A female specimen of Polydesmus denticulatus was identified by comparison of COI sequences, which shows the value of complementing traditional morphology with molecular systematics. Analysed COI sequences of individuals from a Serbian Cave and from the Abaligeti Cave showed only 0.79% difference, which may suggests that the originally epigean species started to colonize the underground habitats and to evolve in isolation during the recent geological past. According to the performed phylogenetic analysis, it was stated that based on the available data, the closest relative of B. troglobius is B. herzogowinensis, a polydesmid species known from the Balkans. Based on the
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available Polydesmida COI sequences, I made attempts for species and genus delimitation. Within the genus Brachydesmus, differences above 4% already meant distinct species, while 11% difference referred distinct genus. 5. Providing new data for estimating the conservation states of the taxa detailed in the thesis. Our knowledge on the distribution of the revised species and subspecies were broadened by new data. Current status of these habitats and the Abaligeti Cave and the Mánfai-kőlyuk Cave – which had been previously studied in zoological aspect – were discussed in detail in the thesis. Referring to their extremely narrow distributional range (< 20 km2) and to the deterioration of the Mánfai-kőlyuk Cave – type locality of several endemic taxa –, N. molnari, N. gebhardti, P. hungarica hungarica and P. hungarica robusta were suggested to be placed into the ’Vulnerable’ IUCN category. Although, the ’Hungarian blind snail’ (B. hungaricum) is already among the ’Vulnerable’ Red List species, in possession of our new results, I recommended to manage the two currently known populations as two separate conservation biological units. Though, the polydesmid B. troglobius is known from some of the caves of Serbia, Slovenia, Croatia and Montenegro, in Hungary it has only two distributional records (Abaligeti Cave and Törökpince Cave), hence, I suggested its legal protection. Furthermore, I made suggestion for a certain method of the removal of show caves’ lamp flora, which does not result in the destruction of its invertebrate inhabitants. 6. Broadening of faunistic data of the Mánfai-kőlyuk Cave and the Abaligeti Cave by 7 and 25 new records apart the known fauna, respectively, as well as contribution to the better knowledge on the subterranean invertebrate diversity of the Western Mecsek Mts. by 12 newly studied caves. Apart from the revised species and subspecies, I have revealed 105 further taxa from the 14 examined caves. I have created the faunal lists of each cave. Although, the majority of the collected species (47%) belonged to the trogloxene category, proportion of subtroglophile (24%) and eutroglophile (26%) was remarkable, which mainly came from the different groups of snails, diplopods, isopods, springtails and coleopterans. Including the 7 taxa detailed in the thesis,
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10% of the species collected in the 14 caves belonged to the troglobiont category. In more than half of the newly studied 12 caves at least one troglobiont taxon occured. Among the newly revealed species of the Mánfaikőlyuk Cave there were some urban habitat-dwelling species, which may refers to the maleficence of the cave’s artificial utilization. The earthworm Helodrilus oculatus found in the Abaligeti Cave and the Spirál Cave was proved to be new for the fauna of Hungary. 7. Contribution to the better knowledge on the fauna of the Abaligeti Cave and the Mánfai-kőlyuk Cave by collecting in previously not studied sampling sites and by application of new collecting methods. I gained valuable data from several sampling sites of the Abaligeti Cave, that, at time of zoological investigations of the first half of the 20th century were partly undiscovered, or were not possible to study in the lack of proper technics and equipments. New sampling sites in the Mánfai-kőlyuk Cave are related to the changed, artificial environment. I successfully applied some collecting methods (e.g. leaf litter traps, epikarstic water collectors), which were previously unprecedented in the two caves.
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4. LITERATURE
ANDRÁSSY, I. (1959): Weitere Nematoden Aus Der Tropfsteinhohle Baradla. (Biospeologica Hungarica V.) Acta Zoologica Academiae Scientiarum Hungaricae 5: 1-6. ANGYAL, D., BALÁZS, G., ZAKŠEK, V., KRÍZSIK, V., FIŠER, C. (2015): Redescription of two subterranean amphipods Niphargus molnari Méhely, 1927 and Niphargus gebhardti Schellenberg, 1934 (Amphipoda, Niphargidae) and their phylogenetic position. ZooKeys 509: 53–85. BAJOMI, D. (1977): Áttekintés a magyarországi barlangok faunájáról. Karszt és Barlang 17 (1-2): 23-28. BOKOR, E. (1921): A magyarhoni barlangok ízeltlábúi. Arthropoden der ungarischen Grotten. Barlangkutatás 9: 1-22. DÓZSA-FARKAS, K., LOKSA, I. (1970): Die systematische Stellung der Palpigraden-Art Eukoenenia austriaca vagvoelgyii (Szalay, 1956) und die bisher bekanntgewordenen Fundorte aus Ungarn. Opuscula Zoologica, Budapest 10(2): 253-261. DUDICH, E. (1932): Biologie der Aggteleker Tropfsteinhöhle ‘Baradla’ in Ungarn. Verlag Speläologisches Institut, Wien, 246 pp. GEBHARDT, A. (1934): Az Abaligeti barlang élővilága. Mathematikai és Temészettudományi Közlemények 37: 1-264. GEBHARDT, A. (1967): A Mecsek hegység állatvilágának térbeli elterjedése élőhelyek szerint. Janus Pannonius Múzeum Évkönyve 12: 7−14. MÉHELY, L. (1924): Protelsonia hungarica, nov. gen., n. sp. Ein blinder Isopode aus Ungarn. Zoologischer Anzeiger 58: 353-357. SOÓS, L. (1927): Adatok a magyarországi barlangok Mollusca-faunájának ismeretéhez. Állattani Közlemények 24: 163-180. ZICSI, A. (1974): Ein neuer Höhlen-Regenwurm (Oligochaeta: Lumbricidae) aus Ungarn. Acta Zoologica Academiae Scientarium Hungaricae 20: 227-232.
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5. LIST OF PUBLICATIONS
Own publications which served as bases of theses ANGYAL, D., BALÁZS, G., ZAKŠEK, V., KRÍZSIK, V., FIŠER, C. (2015): Redescription of two subterranean amphipods Niphargus molnari Méhely, 1927 and Niphargus gebhardti Schellenberg, 1934 (Amphipoda, Niphargidae) and their phylogenetic position. ZooKeys 509: 53–85. (Impact Factor: 0,933) ANGYAL, D., BALÁZS, G. (2013a): Distinguishing characters of Niphargus gebhardti Schellenberg, 1934 and Niphargus molnari Méhely, 1927 (Crustacea: Amphipoda): a clarification. Opuscula Zoologica, Budapest 44(1): 3-8. ANGYAL, D., KORSÓS, Z. (2013): Millipedes (Diplopoda) of twelve caves in Western Mecsek, Southwest Hungary. Opuscula Zoologica, Budapest 44(2): 99-106. ANGYAL, D. (2012a): Újabb adatok a magyar vakcsiga (Bythiospeum hungaricum (Soós, 1927); Gastropoda, Hydrobiidae) elterjedéséről az élőhelyében bekövetkezett változások tükrében. Állattani Közlemények 97(2): 163-170. ANGYAL, D. (2012b): A Mánfai-kőlyuk gerinctelen faunájának alakulása a vízmű általi hasznosítás tükrében - előzetes eredmények. Természetvédelmi Közlemények 18: 24-33.
Further own publications cited in the PhD thesis
BALÁZS, G., ANGYAL, D., KONDOROSY, E. (2015): Niphargus (Crustacea: Amphipoda) species in Hungary: literature review, current taxonomy and the updated distribution of valid taxa. Zootaxa 3974(3): 361-376. (Impact Factor: 0,906)
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HORNOK, S., KONTSCHÁN, J., KOVÁCS D., ANGYAL, D., GÖRFÖL, T., POLACSEK, ZS., KALAMÁR, ZS., D. MIHALCA, A. (2014): Bat ticks revisited: Ixodes ariadnae sp. nov. and allopatric genotypes of I. vespertilionis in caves of Hungary. Parasites & Vectors 7:202: 1-9. (Impact Factor: 3,43) SALAMON, G., DÁNYI, L., ANGYAL, D., BALÁZS, G., FORRÓ, L. (2014): A Baradla gerinctelen faunája. In: Gruber P. & Gaál L. (eds): A Baradla-Domicabarlangrendszer - A barlang, mely összeköt. Aggteleki Nemzeti Park Igazgatóság, Jósvafő, 2014. ISBN 978-615-80050-0-5, pp. 279-306. SZEDERJESI, T., ANGYAL, D., BALÁZS, G., DÁNYI, L. (2014): Remarks on the earthworm genus Helodrilus Hoffmeister, 1845 with new epigean and subterranean records (Oligochaeta, Lumbricidae). Opuscula Zoologica, Budapest 45(2): 181-188. MOCK, A., TAJOVSKÝ, K., ŽUROVCOVÁ, M., ANGYAL, D., KOCOUREK, P. (2014): Hungarosoma bokori Verhoeff, 1928 (Diplopoda, Chordeumatida), a tiny and enigmatic millipede: Redescription and new light to its systematics, ecology and biogeography. In: Tuf, I. H. & Tajovský, K (eds): Book of Abstracts, 16th International Congress of Myriapodology, 20-25 July 2014, Olomouc, Czech Republic, p. 58. ANGYAL, D., KRÍZSIK, V., FEHÉR, Z. (2013): Magyar vakcsiga (Bythiospeum hungaricum (Soós, 1927)) egy szisztematikai probléma vizsgálata genetikai módszerekkel. © http//: www.mecsek.gportal.hu, 1-5. ANGYAL, D., BALÁZS, G. (2013b): New data to the distribution of four aquatic troglobiont macroinvertebrate species in some caves of the Mecsek Mountains (SW Hungary). In: Filippi, H. & Bosák, P. (eds): 16th International Congress of Speleology, 21th to 28th July, 2013. Brno, Czech Republic, Proceedings, Volume 2. Czech Speleological Society, Praha, pp. 426-429. BALÁZS, G., ANGYAL, D. (2013): A magyarországi vakbolharákfajok (Amphipoda: Niphargus spp.) értékelő irodalmi áttekintése. Állattani Közlemények 98(1-2): 111-119.
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Other publications/conference summaries related to the topic ANGYAL D., JAKSCH K., BAMBERGER S. & FEHÉR Z. (2015): Adventures in the field and in the lab – investigation of an enigmatic door-snail species, Montenergrina apfelbecki (Sturany, 1907). American Conchologist 43(3): 24-30. BALÁZS, G. & ANGYAL, D. (2015): Study on Niphargus aggtelekiensis Dudich, 1932 (Crustacea: Amphipoda) populations in the caves of the Aggtelek Karst. (10th Scientific Conference „Research, Use and Protection of Caves”, 22-25 September 2015, Roznava (SK) – Bódvaszilas (HU)) Aragonit 20(1): p. 68. ANGYAL, D., BALÁZS, G. (2014): Overview of the Hungarian Niphargus (Crustacea: Amphipoda) species. In: Palacios Vargas J. G. (ed): Abstract Book, 22st International Conference on Subterranean Biology, 31 August5 September 2014, Juriquilla, Querétaro, México, p 31. MOCK A., KOVÁČ L., PAPÁČ V., ĽUPTÁČIK, P., PARIMUCHOVÁ, A., ANGYAL, D., KOŠEL, V. (2014): Suchozemské bezstavovce jaskynného systému Domica-Baradla. – In: MANKO P. & BARANOVÁ B. (eds): „Zoológia 2014“, 19. Feriancove dni, 20. – 22. november 2014, Prešovská univerzita v Prešove pp. 144-146. ANGYAL D., KANCSAL B. (2011): Gerinctelen faunisztikai vizsgálatok a Mecsek hegység barlangjaiban. XVII. Ifjúsági Tudományos Fórum, 2011. április 21., Keszthely .ISBN 978-9639-42-3, 11 pp.
Related oral and poster presentations DÁNYI L., BALÁZS G., ANGYAL D. (2015): Mint giliszta a vízben – avagy Zicsi András nyomdokán a Baradla Rövid-Alsó-barlangban. A Magyar Biológiai Társulat Állattani Szakosztályának 1028. előadóülése, 2015. december 2, Budapest
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A. MOCK, Ľ. KOVÁČ, V. PAPÁČ, P. ĽUPTÁČIK, A. PARIMUCHOVÁ, L. FORRÓ, D. ANGYAL, L. DÁNYI, G. SZÉL, G. BALÁZS, V. KOŃEL, P. FENĎA (2015): Terrestrial and aquatic arthropods (Arthropoda) of the Domica-Baradla cave system. Az Aggteleki- és Szlovák Karszt Kutatása és Védelme Tudományos Konferencia, 2015. szeptember 24, Bódvaszilas ANGYAL D., DÁNYI L. (2015): Állatok az örök éjszakában - barlangzoológiai kutatások Magyarországon. Kutatók Éjszakája, Magyar Természettudományi Múzeum, 2015. szeptember 25, Budapest G. BALÁZS, D. ANGYAL (2015): Study on Niphargus aggtelekiensis Dudich, 1932 (Crustacea: Amphipoda) populations in the caves of the Aggtelek Karst. Az Aggteleki- és Szlovák Karszt Kutatása és Védelme Tudományos Konferencia, 2015. szeptember 24, Bódvaszilas
ANGYAL D. (2014): Kis barlangzoológia – újabb kutatások a hazai barlangokban. „A Mecsek barlangjai” konferencia, 2014. június 20, Abaliget ANGYAL D., BALÁZS G., DÁNYI L. (2014): A hazai barlangzoológia múltja és jelene. XVIII. Karsztfejlődés Konferencia, 2014. május 30, SzombathelyBük BALÁZS G., ANGYAL D. (2014): Vakbolharák ki-kicsoda a magyar faunában. A Magyar Biológiai Társaság Állattani Szakosztályának 1014. előadóülése, 2014. február 5, Budapest BALÁZS G., ANGYAL D. (2013): Chase in history after the endemic Niphargus (Crustacea: Amphipoda) species of Hungary. 16th International Congress of Speleology, 25th July, 2013, Brno, Czech Republic ANGYAL D. (2013): A mecseki barlangok gerinctelen faunája – újabb kutatások. Barlangok Napja, 2013. június 22, Abaliget
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BALÁZS G., ANGYAL D. (2013): A magyarországi Niphargus fajok kiértékelő áttekintése. X. Makroszkopikus Vízi Gerinctelenek Kutatási Konferencia, 2013. 04. 13, Szalafő ANGYAL D., BALÁZS G. (2013): A Niphargus gebhardti Schellenberg, 1934 és a Niphargus molnari Méhely, 1927 (Crustacea: Amphipoda) morfológiai elkülönítő bélyegeinek bemutatása. X. Makroszkopikus Vízi Gerinctelenek Kutatási Konferencia, 2013. 04. 12, Szalafő ANGYAL D., BALÁZS G. (2012): Niphargusok nyomában – a hazai vakbolharák fauna jelenlegi kutatása. Barlangkutatók Szakmai Találkozója, 2012. november 18, Pécs ANGYAL D. (2012): Újabb adatok három troglobiont vízi makrogerinctelen taxon elterjedéséről a Mecsek hegység barlangjaiban. IX. Makroszkopikus Vízi Gerinctelenek Kutatási Konferencia, 2012. április 12, Gyula ANGYAL D., KRÍZSIK V., FEHÉR Z. (2012): Molekuláris taxonómiai vizsgálatok a magyar vakcsiga (Bythiospeum hungaricum (Soós 1927)) két mecseki barlangból származó egyedein. IX. Makroszkopikus Vízi Gerinctelenek Kutatási Konferencia, 2012. április 12, Gyula ANGYAL D. (2012): A mecseki barlangok gerinctelen faunájának diverzitása – újabb adatok. A Magyar Biológiai Társaság Állattani Szakosztályának 999. előadóülése, 2012. március 7, Budapest ANGYAL D. (2011): Gerinctelen faunisztikai vizsgálatok a Mecsek hegység barlangjaiban. Barlangkutatók Szakmai Találkozója, 2011. november 12, Székesfehérvár ANGYAL, D. (2011): Gerinctelen faunisztikai vizsgálatok a Mánfai-kőlyukban – egy sokat megélt barlang múltja, jelene és jövője. VII. Természetvédelmi Biológiai Konferencia, 2011. november 5, Debrecen
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