The History of Cytogenetic Investigations of Spiders in Turkey

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Kumbicak Z* Review Article Volume 3 Issue 4 Department of Molecular Biology and Genetics, Nevşehir Hacı Bektaş Veli University, Turkey *Corresponding author: Received Date: Published Date: June 12, 2020 Zubeyde Kumbicak, Department of Molecular Biology and Genetics, July 06, 2020 Faculty of Art and Science, Nevsehir Hacı Bektas Veli University, 50300, Nevşehir, Turkey, DOI: 10.23880/izab-16000232 E-mail: [email protected] Abstract

Spiders are an ancient group of animals that probably originated in the Devonian period about 400 million years ago. Over time, they have adapted to major changes in climate and fauna and have become widespread on all continents, living in almost any terrestrial habitat. It is known that approximately 50000 spider species belonging to 120 families worldwide, today. and this number increases almost every day due to new records are being given from many countries. Despite the high diversity of species, the number of cytogenetic studies on spiders is very scarce. Upto now, 868 species belonging to the 74 families were investigated by cytogenetically. Although more than a thousand species of spiders in Turkey, the Cytogenetic studies are insufficient. Thus, spiders are the most richest group of arachnids among arthropods. In this study cytogenetic studies on Keywords:spiders in Turkey were reviewed.

Convergent Evolution; Rheumatoid Arthritis; Typhoid; HIV Abbreviations: IntroductionSCS: Sex Chromosome System. pedipalpus, which has a bulging structure in male spiders, is an important structure in the transfer of sperm to the female. Basic Characteristics of Spiders The remaning of the four extremity are walking legs. The opistosoma is the part of the body that contains the excretory system, reproductive organs and knitted glands. Male spiders have a pair of testes on either side of the abdomen. The sperm Spiders are one of the richest group in Chelicerata. channels are long and curved, and the matured sperm are The animals in this group do not have antennas [1]. The ejected from the middle of the epigastric slit through these body structure of spiders consists of two main parts called channels. Sperm expelled from this opening is stored in the cephalothorax (prosoma) and abdomen (opistosoma). These embolus at the ends of the pedipalps of male spiders, and two parts are connected to each other by a narrow part called from there, sperm is transferred to the female during mating pedicel. The dorsal part of cephalothorax is covered with a [3,4]. arestructure located called ö carapace, and its ventral part is covered with a plate called the sternum. The mouth, eyes and extremities All spiders are carnivores. Some of them live freely, while n the cephalothorax. others live dependent on the net they built. Ecological and faunistic researches in forests, fields, gardens and grasslands Generally, spiders have simple eyes and the number of has shown that spiders are the most common predators. A eyes varies between 2-8. In determining family, characters large number of spiders live in soil layers, stony, rocky and related to the position, number and structure of the eyes are woodland, in the remains of organic matter and in terrestrial used. There are 6 pairs of extremities in the cephalothorax. habitats such as plant tissues. A small number of spiders Among them, the first extremity is called chelicer located live on the water shores, on the surface of the freshwater on the front of the mouth with poison glands in their base and inside them. Since they are carnivorous, most of their [2]. The second pair of extremity is called pedipalpus. It is nutrients are insects [5]. responsible for keeping food during feeding. In addition, The History of Cytogenetic Investigations of Spiders in Turkey Int J Zoo Animal Biol 2 International Journal of Zoology and Animal Biology

Cytogenetics of Spiders

genera and 90 species) and Lycosidae (15 genera and 87 species) [16]. Although some species are endemic, some are The phase in which chromosome morphology is best very widespread throughout the country. The first study of observed during cell division is the metaphase. At this the spider cytogenetics in Turkey was carried out by Akan, et stage, chromosomes can be seen by regular condensation al. [17]. The authors were studied of three species belonging of chromatin threads. Karyotype refers to the ordering of to the families Lycosidae Arctosa and Theraphosidae. perita Adult and chromosomes in a cell according to a certain order after Lycosasubadult narborensis individuals were used and as a resultChaetopelma the diploid matching, based on the chromosome morphology, number anatolicumchromosome number of (Latreille 1799), and size of an individual or species [6]. Karyotype features (Walkenaer 1806) and such as diploid number (2n), sex chromosome system (SCS) (Schmidt, Smith 1995) was 2n=12, 2n=18 and chromosome morphology are important factors in and 2n=16, respectively. There was no data about the sex understanding the cytogenetic structure of spiders. chromosome system of the spiders studied.

Cytogenetic studies of spiders were firstly made by In 2009, the karyotype of four species of the Gnaphosidae ♂ 1 2 Callilepis cretica Carnoy [7], using gonads of male or females that imbedded were raported Drassodes by Kumbıcak, pubescens et al., Drassyllus [18]. The pumilus karyotype in paraffin, sectioned, and stained with Heidenhain’s iron formula was 2n Zelotes=22 (X strandiX 0) for all species, haematoxylin. Visualisation and evaluation of chromosomes (Roewer 1928), (C L were not easy by the protocol used. Thus decades later Koch 1839) and (Nosek 1905). Sharma, et al. [8] and Becak, et al. [9] were firstly obtained chromosomes using the squash methods of aceto-orcein and The diploid chromosome number and the♂ sex aceto-carmin [10]. After that colchicine to promote solution an was increase used for in chromosome system of five species belonging to the families 1 2 ♂ 1 2 karyological preparations of spider gonads for both males Gnaphosidae Nomisia and Lycosidae conigera were (Spassky determined 1941), Haplodrassus as 2n = 22 and females by Pinter, et al. [11] morosus(X X 0) and (O. 2n Pickard-Cambridge = 28 (X X 0), respectively 1872), and Haplodrassus the studied the number of cells in mitotic and/or meiotic metaphase. In dalmatensisspecies were (L Pardosa bifasciata (CL Koch 1834) the same decade Brum Zorrila, et al. [12] were used a fixative and Arctosa cinerea solution (3:1; methanol:acetic acid) and Giemsa solution Koch 1866), as a stain. In 1977 Matsumoto, et al. [13] launched the use [19].Zelotes Taşdemir, petrensis et al. [20] were reported of observation of chromosomes in spider embryos which Zelotesthe karyotype aeneus characteristics of three speciesNomisia belonging conigera to are valuable source of mitotic metaphase cells due to the the Gnaphosidae as (CL Koch 1839)Theridion 2n=11, high rate of cellular division that occurs during embryonic pictum (Simon 1878) 2n=9 andSteotoda triangulosa development. After that Wang, et al. [14] were described (Spassky 1941) 2n=10; two species of Theridiidae, the number of tissues that can be used in spider cytogenetic Trochosa (Walckenaer ruricula 1802) 2n=14 and studies, such as cerebral ganglion, cultured blood cells and (Walckenaer 1802) 2n=12; one species for Lycosidae, gonads. Among them, the gonads were usablecomparing Karyotype features (De Geer of Drassyllus 1778), 2n=9. praeficus than the other tissues for cytogenetical analysis [10]. Thanatus imbecillus (L. Koch 1866) There are many different sex chromosome systems (Gnaphosidae) and (Philodromidae) (L. among spider species. Most for of themmany havespecies, more and than for one male X Koch 1878) were obtained using chromosomes from gonads. 1 2 system of D. praeficus and T. imbecillus ♂ 1 2 chromosome in their karyotypes. consists of two different The diploid chromosome number and the sex chromosome and 2n♂ 1 2 X chromosomes, X and X was 2n = 22 (X X 0) spiders, this system is considered to be ancestral. This sex Berinda= 28 (X Xhakani0), respectively [21]. In the year 2014,♂ =22 a chromosome system is generally seen in primitive spider total of 23 species were investigated by cytogenetically as 1 2 1 2 Berinda ensigera groups namely Mesothelidae and Mygalomorphae. TheY other and follows: Chatzaki & Seyyar 2010 2n 1 2 1 2 1 2 1 2 1 2 2n ♂ 1 2 Trachyzelotes lyonneti examples of sex chromosome systems of spiders are X X 0, (X X 0), (O. Pickard-Cambridge 1874) 3 3 4 3 3 4 5 2n ♂ 1 2 Trachyzelotes malkini X0, X X X 0, X X Y, X X X X Y, X X X Y and X X X X X =22 (X X 0), (Audouin, 1826) ♂ 1 2 Zelotes caucasius XnYn0. “0” in the sex chromosome systems indicates that =22 (X X 0), Platnick & Murphy 2n♂ = 1 2 Thanatus pictus there is no Y chromosome [15]. 1984 2n =22 (X X 0), (L. Koch 1866) ♂ = 1 2 Tibellus macellus ♂ 22 (X X 0) (Gnaphosidae); L. Koch 1 2 Neon reticulatus In Turkey, a total of 1117 spider species (with two 1881 2n ♂28 (X X 0), PeucetiaSimon 1875virescens 2n =24 subspecies) belonging to 52 families have been found. Among (X X 0) (Philodromidae); (Blackwall ♂ = 1 2 them, the richest families are Gnaphosidae (30 genera 1853) 2n =21 (X0) (Salticidae); (O. and Loxosceles rufescens ♂ = 21 (X1 2 and 145 species), Salticidae (42 genera and 143 species), Pickard-Cambridge 1872) 2n 28 (X X 0) (Oxyopidae) Linyphiidae (66 genera and 128 species), Thomisidae (14 (Dufour 1820) 2n X Y)

Pterotricha kochi References Pterotricha lesserti ♂ = 1 2 (Sicariidae) [22]; Drassyllus sur (O. Pickard-Cambridge 1872) and Dalmas 1921 2n 22 (X X 0) 2n♂ 1 2 Nomisia exornata ♂ (Gnaphosidae) [23]; Tuneva & Esyunin 2003, 1. Hickman CP, Roberts LS, Keen SL, Eisenhour DJ, Larson 1 2 Nomisia orientalis ♂ 1 2 =22 (X X 0), (C. L. Koch, 1839) 2n =22 A, et al. (2016) Zoology Integrated Principles. In: Gündüz Sitticus caricis ♂ 1 2 Xysticus (XgallicusX 0), and ♂ Dalmas 1921Pax 2n islamita=22 (X X 0), E, et al. (Eds.), Palme Publishing, Kültür pp: 35-823. Publisher, Niğde. (Westring 1861) 2n =28 (X X 0); ♂ 1 2 Drassodes lutescens: 2n♂ rd Simon 1875 2n =23, X0 and in (Simon 2. Babaşoğlu A (1999) Arachnida. Micaria albovittata 2n♂ 1 2 Cheiracanthium 1873) 2n =42, (X X 0) [24]; =21 mildei 2n♂ 1 2 Cheiracanthium pennyi: 2n♂ 3. Foelix RF (2011) Biology of Spiders. 3 (Edn.), Oxford (X0), =22 (X X 0), 1 2 Philodromus lividus: 2n♂ 1 2 University press, Newyork. =26 (X X 0), =26 (X X 0) and =28 (X TegenariaX 0) [25]. elysii 4. Seyyar O (2009) Doğu Akdeniz Bölgesi’nin Yer

♂ 1 2 Örümcekleri (Araneae, Gnaphosidae) Faunası. Science In 2018, the karyotype formula of Hersiliola bayrami Danisman ♂ 1 2 Institute, Erciyes University, PhD Thesis, Kayseri. Brignoli, 1978 were found as 2n =42 (X X 0) [26,27], and properties of Dysdera crocata 3 Steatoda, et al. as grossa 2n =35 (X X X 0). The 5. Alazzouzi SFK (2020) Characterizatıon of morphological next year, karyotype features based on diploid number and CL Koch, 1838 spider ♂ 1 2 sex chromosomeEratigena system agrestris of (C.L. Koch 1838) species with scanning electron microscope (Arachnida: was reported as 2n =22 (X X 0) for Turkish populations Araneae). Science Institute, Kastamonu University, [28], for (Walckenaer 1802) the diploid Master Thesis, Kastamonu, Turkey. 2n♂ 1 2 Gnaphosa lugubris chromosome number and sex chromosome system was Chondrostoma beysehirense ♂ 1 2 =42 (X X 0) [29], and for (CL Koch, 6. Gündoğdu H (2016) Cytogenetic studies on Endemic Beyşehir Karaburun Fish, 1839) the karyotypeDrassodes formula bifidus was 2n =22 (X X 0) [30]. The (Bogutskaya, 1997). Science Institute, Selçuk University, andlast Drassodes study on spiderserratichelis cytogenetics (Roewer was made by Kumbıçak, Master Thesis, Konya. et al. [31] on (Kovblyuk and Seyyar 2009)♂ 1928), according to the 1 2 D. bifidus and D. serratichelis 7. Carnoy JB (1885) La cytodiérese chez les arthropodes. results the karyotype characteristics of them was 2n =22 La Cellule 1: 191-440. (X XConclusion0) for both . 8. complementSharma GP, Jandeand meiosis SS, Tandon in Selenops KK (1959) radiatus Cytological Latreille studies on the IndianLeucauge spiders decorata IV. Chromosome

There are 1117 spider species known in Turkey, (Selenopidae) and (Blackwall) among them almost 60 species (5.37%) were investigated (Tetragnathidae) with special reference to XXX0-type by cytogenetically. belonging to the families Agelenidae, of male sex determining mechanism. Research Bulletin Gnaphosidae, Hersiliidae, Lycosidae, Oxyopidae, (N.S.) of the Panjab University 10(1): 73-80. Philodromidae, Salticidae, Sicariidae, Theraphosidae, Theridiidae, and Thomisidae. Of these families, Gnaphosidae 9. Becak W, Becak ML (1960) Constituição cromossômica is the most studied group due to its widespread. The studies de duas espécies de aranhas do gênero “Loxosceles”. have provided information about the diploid chromosome Revista Brasileira de Biologia 20(4): 425-427. number and sex chromosome systems of the species, but chromosome banding studies have not been included. 10. Araujo D, Schneider MC, Paula-Neto E, Cella DM (2012) According to these studies, cytogenetic features were Sex chromosomes and meiosis in spiders: a review. In: generally conservative at the family. Swan A (Eds.), Meiosis – Molecular Mechanisms and Cytogenetic Diversity, InTech, Croatia, Rijeka, pp: 87- The reproduction period of spiders takes place in a 108. short period of time, which reduces the rate of obtaining chromosomes from them. Thus, spider cytogenetic studies 11. Pinter LJ, Walters DM (1971) Karyological studies I. A are progressing slowly. As a result, since there are many study of the chromosome numbers and sex-determining families that have not been studied yet, samples of these mechanism of three species of the genus Phidippus families should also be investigated. Therefore, more field (Aranea: Salticidae, Dendryphantinae). Cytologia 36(1): studies should be organized and more samples should be 183-189. collected and applied. Moreover, chromosome banding 12. Brum Zorrilla N, Cazenave AM (1974) Heterochromatin techniques such as C-banding and NOR staining should be localization in the chromosomes of Lycosa malitiosa included in the studies to obtain detailed data. (Arachnida). Experientia 30(1): 94-95.

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