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Biology of the Saw-Legged Bush Crickets (Saga Spp .) Focusing On

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Biology of the Saw-Legged Bush Crickets (Saga Spp .) Focusing On Biology of the saw-legged bush crickets (Saga spp.) focusing on Saga pedo (PALLAS, 1771) Outline of the PhD thesis Kolics, Balázs Supervisors: Kondorosy, Előd, CSc Müller, Tamás, PhD & Pannon University Szent István University Pannon University, Georgikon Faculty Keszthely, 2009 1. Introduction, aim of study Conservation biology is a significant field of biological research, because of the accelerating distinction of species mainly because of human impacts. Saginae species are amongst the largest insects of Eurasia. Their populations are attracted to xerotherm habitats that they found on rocky steppes of middle mountains. They live in isolated sparse populations being of low density especially in imago stage. Being flightless, Saga species are less able to move from one habitat to another. As particular orthopterans, these katydids are obligatory insectivores being on the top of the arthropod food pyramid in their habitat. Despite being rare, Saga species are not protected, except one representative of the genus. The matriarchal katydid, Saga pedo Pallas 1771 is the only species of its genus living in Hungary as well, in addition being one of our the largest insect in our fauna. Saga pedo is the only tetraploid species of its genus - comprising 13 species - reproducing in a parthenogenic way. 2 Its distribution area covers a territory several fold larger than that of its congeners. Conservational value in Hungary is 50.000Ft. The aim of my study was to comply an up-to date habitat list of the matriarchal katydid, and possibly to discover new populations. In phenological measurements I wanted to determine the number of larval stadia of S. pedo undergoes in its development - on the reason of contradiction in literature - included parameters describing each stadium. In egg measurements I wanted to deal with ootaxonomic parameters such as length, diameter, weight and number of micropyles of the egg of European species have not been described yet. In sonic examination the aim was to describe song of Saga species, yet unknown and a comparison of the song of the two subspecies, S. campbelli campbelli and S. campbelli gracilis. One of the main aim of my dissertation was to uncover natural relationship in genus Saga, complementing results with morphometry where it was necessary, and also with bioacoustic information effectively applicable for solving taxonomic problems in 3 orthopterans. Another point of the genetic examination was to test the hybrid status of the eggs deriving from a copulation between the matriarchal katydid and its close congener S. c. campbelli. Furthermore, I wanted to eliminate the initial and the terminal diapause of the long developing species S. pedo and S. rammei, in order to base the reintroducing them to extinct habitats. 2. Materials and methods (1) Compilation of the habitat list of the matriarchal katydid was based on literature, personal contacts and a homepage established for this purpose. Furthermore, populations were checked via field trips as well. (2) Larval stadia was determined by measuring length of hind femur and ovipositor. Altogether 156 S. pedo specimen were involved into the examinations. They originated from the Nagy, Barnabás’s (PhD, MTA-NKI) and own breeding. Ootaxonomic measurements were carried out on species S. pedo, S. natoliae, S. rammei, 4 and S. c. campbelli; sample examined amounted to 580 eggs. Recorded were length, diameter, weight and number of micropyles. (3) In bioacoustic examinations, 15 specimen of the following European species were involved: S. natoliae, S. rammei, S. hellenica, S. c. campbelli, S. c. gracilis. Sound tracks were recorded at the Georgikon Faculty and the Hungarian National History Museum, and were visualized with Adobe Audition 1.5 software by oscillographic, sonographic and spectrographic examinations. (4) Altogether, 27 specimen were involved in phylogenetic studies, from the following species: S. pedo, S. rammei, S. natoliae, S. hellenica, S. c. campbelli, S. c. gracilis, S. cappadocica, and S. e. ephippigera. Three mitochondrial (CO I, Cyt b, 16S) and one nuclear (ITS2) markers were used in sequencing. Amplified fragments were of a length in total 2453-2461 bp. Analyses were carried out with Maximum parsimony and Bayesian inference in four phylogenic tree. Results were evaluated together with those derived from bioacoustics and 5 morphometry (principal component analysis) in the case of S. c. campbelli, S. c. gracilis and S. pedo, in order to get whole scale information to solve taxonomic problems of the two subspecies. (5) The matriarchal katydid was paired with a male specimen of its congener, S. c. campbelli. Eggs derived from this copulation were treated with heat shock in order to gain whole embryos, thus appropriate amount of DNA for testing the hybrid status of the offspring (n=2 eggs with micropyles). FluoMEP analysis was used to detect the presence or absence of paternal DNA in the F1. In FluoMEP, 26 potentially informative combination of common fluorescent (labeled) and RAPD (unlabelled) primers were used for testing the potential hybrid and the control (without micropyles) eggs. (6) In additional examinations, embryonic development of S. pedo and S. rammei was investigated and traced by measuring the diameter of the egg. Elimination of the initial diapause was carried out according to literature data. For the investigation of the alternative distribution of the matriarchal katydid, were used. Eggs were 6 embedded into the maternal animal or into abdomen of Acheta domesticus and were fed with Falco tinnunculus and Lacerta viridis, as model species . 3.Results and discussion (1) An UTM distribution map of S. pedo containing 100 habitat was complied in the dissertation. It contains 29 more populations comparing to the latest study compiled by Bauer et al. (2002). Furthermore, one habitat of the matriarchal katydid and one of S. hellenica were transmitted in the dissertation. (2) According to the specimen examined the matriarchal katydid always goes through six larval stadia until moulting to imago. It showed a result different to previous authors, where 8-9 (Bérenguier 1907) or 5-6 (Schall 2002) stadia was transmitted. In my investigations, each larval stadium was supported by confirming morphological parameters (length of hind femur and ovipositor). According to the ootaxonomic results, no significant differences was observed in the 7 length of the eggs between S. pedo and S. natoliae (p <0,05) . Eggs of S. c. campbelli and S. rammei were however significantly smaller from those of the former two species. The species concerned are significantly different and distinguishable according to weight and the number of micropyles on the egg. For S. rammei and S. c. campbelli ootaxonomic characteristics were firstly given in the dissertation, while those of S. pedo were refined. The highest variability was observed in the eggs of the matriarchal katydid according to the length, diameter and the number of micropyles (mp). The latter distributed between 0 and 6 mp/egg. To sum up, 1 mp/egg was found to be the most frequent (27%). The eggs did not possess any micropyles in 52,98% of all eggs (n=151) of S. pedo, while a high variability was found amongst the individuals as well; a specimen was observed with a lack of mp of 90% amongst its eggs layed. The rate of the eggs without micropyles is different (with higher number of samples) to those found by former authors (27%. Sänger and Helfert (1994),, moreover, individual variability was observed. Because of the lack of 8 micropyles, penetrating the sperm into the egg is not possible into many of the eggs. (3) In sonic investigations of S. rammei, I observed the following. The calling song is long echeme sequence in which the echeme repetition period was found to be 1,3- 1,7 sec on 23-26°C. Apart from the first few syllables, an echeme consists of one type of syllables, with a repetition of 40-48 syllable/s. In the initial part (about one third) of the echeme, syllables become successively louder and longer composed of more impulses. .After the initial crescending part syllable amplitude reaches its maximum and becomes constant during the main part of the echeme. The sonographic and oscillographic pattern of the echemes showed that the signal is mainly amplitude modulated: Syllables are composed of two impulse- series: a quieter and a louder one. For most part of the echeme the quiet serie is shorter and composed of a lower number of impulses than the louder one. Furthermore, during the main part of the echeme the component impulses follow each other with an increasing repetition rate and decreasing amplitude. For the louder series. the 9 opponent pattern is typical. Detailed pattern of the calling song of S. rammei was transmitted in oscillograms, spectrograms and sonograms. The distribution area of S. rammei is the smallest comparing to the congeners, thus a description of the song of this animal can ease it detection with a high reliability. The four taxa examined were found to be clearly different according to four rhythmic parameters. Interestingly it was so in the case of S. c. campbelli and S. c. gracilis as well, that was supported by PCA analysis as well. (4) According to molecular markers, Saga species proved to be of monophyletic origin. The European species derive from the Asians. S. ephippigera and S. ornata possess two types of ITS2 sequences, showing ancestral variability. Furthermore ITS2 comprised and insertion of 115 bp. The latter was also found in S. cappadocica, a geographic transitional species. 10 Species groups raised by the results are demonstrated by Table 1. In comparison with those revealed former authors showing whole scale differences. Table 1. Comparative data on the phylogenetic relationship of the Saga species Bold: Most basal species Underlined: origin species of S. pedo The two subspecies and S. pedo shared the same topology both with the protein coding mitochondrial 11 (COXI, Cytb) and the nuclear (ITS2) gene as follows: [(S.
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