{"title":"Morphological variation between Hyla arborea and Hyla orientalis from Bulgaria","authors":"Simeon Lukanov , Irena Atanasova , Emiliya Vacheva , Angel Dyugmedzhiev , Simeon Borissov , Borislav Naumov","doi":"10.1016/j.jcz.2025.09.010","DOIUrl":"10.1016/j.jcz.2025.09.010","url":null,"abstract":"<div><div>The species status of <em>Hyla arborea</em> and <em>H. orientalis</em> is well supported by genetic data, however, there is still a lack of studies that could differentiate between these taxa based on other characteristics. During the mating season in the period 2021–2024, we measured a total of 196 tree frogs (144 male and 52 female) from six sites – three within the range of <em>H. arborea</em> and three within the range of <em>H. orienatalis</em> in Bulgaria. All frogs were anesthetised, measured, photographed and released at the site of capture after they were fully recovered. We assessed eight morphometric traits and corresponding indices, and used statistical comparisons to examine species- and population-level variation. While some indices showed significant differences, results did not support clear-cut morphological separation between species. Notably, one population (Zaychino), genetically identified as <em>H. orientalis</em>, grouped morphometrically with <em>H. arborea</em>. Our results suggest that interspecific morphological differences are subtle and may be more influenced by local environmental adaptation or historical introgression. Although not sufficient for a direct species identification in the field, the results contribute towards a better understanding of the morphological differences between these two congeneric species.</div></div>","PeriodicalId":49332,"journal":{"name":"Zoologischer Anzeiger","volume":"319 ","pages":"Pages 161-172"},"PeriodicalIF":1.5,"publicationDate":"2025-09-19","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"145158663","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":3,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Marcos J.M. Dubeux , Gisele M.N. Vieira Dubeux , Jessika M.M. Neves , Fernanda P. Werneck , Miguel T. Rodrigues , Tamí Mott , Pedro M.S. Nunes
{"title":"Uncovering neotropical rock geckos’ diversity: multiple data sources on an integrated taxonomic approach reveal a new species of Phyllopezus Peters, 1878 (Squamata: Phyllodactylidae) from northeastern Brazilian Atlantic Forest","authors":"Marcos J.M. Dubeux , Gisele M.N. Vieira Dubeux , Jessika M.M. Neves , Fernanda P. Werneck , Miguel T. Rodrigues , Tamí Mott , Pedro M.S. Nunes","doi":"10.1016/j.jcz.2025.09.011","DOIUrl":"10.1016/j.jcz.2025.09.011","url":null,"abstract":"<div><div>Using an integrated taxonomic approach based on a robust data set including external morphology, cranial anatomy, hemipenial morphology, scale microstructure, and molecular data, we describe a new species of gecko of the genus <em>Phyllopezus</em> from mountains in the northeastern Brazilian Atlantic Forest. The species is currently known to occur in <em>Serra da Jiboia</em>, municipality of Elísio Medrado, Bahia state, and <em>Serra de Itabaiana</em>, municipality of Areia Branca, Sergipe state. The new species is genetically related and more morphologically similar to <em>Phyllopezus diamantino</em> and <em>Phyllopezus selmae</em>, sharing all diagnostic characteristics for both these species, but distinguished from their congeners by meristic and morphometric characters. The new species is the first Phyllodactylidae to have its hemipenial morphology described and together with the cranial anatomy and the description of the microstructures of the dorsal scales provided here, it encourages the use of new approaches for the description of the cryptic species present in this species complex.</div></div>","PeriodicalId":49332,"journal":{"name":"Zoologischer Anzeiger","volume":"319 ","pages":"Pages 173-195"},"PeriodicalIF":1.5,"publicationDate":"2025-09-19","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"145158662","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":3,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
{"title":"Description of new species of Oligosita Walker (Hymenoptera: Trichogrammatidae) from India","authors":"Mohsin Ikram , Mohd Yousuf , Mohammad Faisal","doi":"10.1016/j.jcz.2025.09.006","DOIUrl":"10.1016/j.jcz.2025.09.006","url":null,"abstract":"<div><div>The genus <em>Oligosita</em> Walker is an important parasitoid, primarily attacking the eggs of hemipterous insect pests. In the present study, <em>Oligosita barabankiensis</em> Ikram & Yousuf, <em>Oligosita hayati</em> Ikram & Yousuf and <em>Oligosita lakhimpurensis</em> Ikram & Yousuf are described as new species, along with an updated key to the Indian species. Additionally, some existing Indian species of <em>Oligosita</em> are diagnosed and illustrated, along with new records from India. These findings enhance the taxonomy of egg parasitoids and highlight the potential role of <em>Oligosita</em> species in biological control, particularly in managing hemipteran pests in agricultural systems.</div></div>","PeriodicalId":49332,"journal":{"name":"Zoologischer Anzeiger","volume":"319 ","pages":"Pages 153-160"},"PeriodicalIF":1.5,"publicationDate":"2025-09-18","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"145118199","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":3,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Kota Okamoto , Teppei Jono , Hitoshi Tanioka , Toyofumi Sueyoshi , Takanori Matsuo , Mamoru Toda
{"title":"Natural hybridization among species of the genus Gekko (Squamata: Gekkonidae) in Japan: the roles of pre- and postmating isolation","authors":"Kota Okamoto , Teppei Jono , Hitoshi Tanioka , Toyofumi Sueyoshi , Takanori Matsuo , Mamoru Toda","doi":"10.1016/j.jcz.2025.09.008","DOIUrl":"10.1016/j.jcz.2025.09.008","url":null,"abstract":"<div><div>Among the eight Japanese species of the genus <em>Gekko</em>, natural hybridization was reported in four pairs at various frequencies. A recent behavioral study suggested that the species-specific courtship calls of the males act as premating isolation for some species of these geckos; however, others lack species-specific calls, which results in hybridizations. No comprehensive study has been conducted to identify a general pattern in the development of isolating barriers in this group. We examined the hybridization status between eight pairs of seven species using population genetic analyses based on microsatellite markers and allozyme data. We then examined the changes in the frequency of hybrids and the proportion of F<sub>1</sub> hybrids against the total number of hybrids along with increasing genetic distances in a phylogenetic framework. The contribution of the courtship call type as a determinant of the presence/absence of hybridization was evaluated by statistical modeling. The results of the genetic analyses revealed hybridizations between two pairs of three species, each having a species-specific call, in addition to previously reported species pairs. Statistical analyses indicated that the frequency of hybrids between species was negatively correlated with genetic distance, and the proportion of F<sub>1</sub> hybrids increased with genetic distance. The results suggest the progressive development of postmating isolation with genetic divergence in this group. The contribution of call type to the presence/absence of hybridization was not fully supported, in part, because of the hybridizations of one of the species exhibiting species-specific calls. A possible reason for the exceptional hybridizations between the species with species-specific calls is discussed.</div></div>","PeriodicalId":49332,"journal":{"name":"Zoologischer Anzeiger","volume":"319 ","pages":"Pages 196-209"},"PeriodicalIF":1.5,"publicationDate":"2025-09-18","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"145158661","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":3,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
{"title":"Coriophagus tigrilepus sp. nov. (Strepsiptera: Halictophagidae: Coriophaginae): A new species and a new record subfamily from China","authors":"Xuhongyi Zheng , Pengxu Mu , Chuan Liu","doi":"10.1016/j.jcz.2025.09.009","DOIUrl":"10.1016/j.jcz.2025.09.009","url":null,"abstract":"<div><div>Halictophagidae is a strepsipteran family that parasitizes Orthoptera, Diptera, Blattodea and Hemiptera. To date, only 2 genera and 12 species have been recorded in China. In this study, we describe <em>Coriophagus tigrilepus</em> sp. nov., based on a male specimen collected from Hainan Province. This discovery represents the first record of both the genus <em>Coriophagus</em> and the subfamily Coriophaginae in China. The new species characterized by unique maxillary palpi, the absence of apical expansion on the forefemur, a slender R2 in the hindwing and a slender aedeagus with a short apical hook. Additionally, we provide an updated checklist of Halictophagidae species documented in China and present a revised key to all <em>Coriophagus</em> species worldwide.</div></div>","PeriodicalId":49332,"journal":{"name":"Zoologischer Anzeiger","volume":"319 ","pages":"Pages 127-133"},"PeriodicalIF":1.5,"publicationDate":"2025-09-17","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"145106062","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":3,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
{"title":"Two new species of Leptopsis Desutter-Grandcolas, 1996 (Orthoptera: Phalangopsidae: Phalangopsinae) from the Brazilian Amazon rainforest with insights from potential distribution models for the genus","authors":"Lianderson Farias Franco , Beatriz Harumi Kondo Oya , Rodrigo Antônio Castro-Souza , Gustavo Costa Tavares","doi":"10.1016/j.jcz.2025.09.007","DOIUrl":"10.1016/j.jcz.2025.09.007","url":null,"abstract":"<div><div><em>Leptopis</em> is a South-American genus with two subgenera and seven valid species: <em>Leptopsis</em> (<em>Leptopsis</em>) Desutter-Grandcolas, 1996, including <em>Leptopsis</em> (<em>Leptopsis</em>) <em>nauta</em> Desutter-Grandcolas, 1992, <em>Leptopsis</em> (<em>Leptopsis</em>) <em>zumun</em> Desutter-Grandcolas, 1992, <em>Leptopsis</em> (<em>Leptopsis</em>) <em>saussurei</em> Desutter-Grandcolas, 1992, <em>Leptopsis</em> (<em>Leptopsis</em>) <em>ecuadori</em> Gorochov, 2006, and <em>Leptopsis</em> (<em>Leptopsis</em>) <em>ucayali</em> Gorochov, 2011; and <em>Leptopsis</em> (<em>Aberracla</em>) Gorochov, 2011, including <em>Leptopsis</em> (<em>Aberracla</em>) <em>chopardi</em> Desutter-Grandcolas, 1992, and <em>Leptopsis</em> (<em>Aberracla</em>) <em>morona</em> Gorochov, 2011. In this work, we describe two new species, <em>Leptopsis</em> (<em>Leptopsis</em>) <em>vader</em> <strong>sp. nov</strong>. and <em>Leptopsis</em> (<em>Leptopsis</em>) <em>silens</em> <strong>sp. nov</strong>, recorded in the state of Pará, Brazil, the first record of the genus in the country, and predict and discuss the distribution of the genus through Species Distribution Models (SDMs). These two new species are easily distinguished from the others by the shape of the phallic complex sclerites, specifically the pseudepiphallic sclerite A. Additionally, the females exhibit differences in copulatory papillae among species for which these structures are known. We also propose synonymizing <em>Mellomima</em> Desutter-Grandcolas, 2020 under <em>Leptopsis</em> (<em>Aberracla</em>), lowering the rank from genus to subgenus, and moving <em>Mellomima guyanensis</em> Desutter-Grandcolas, 2020 to the subgenus <em>Leptopsis</em> (<em>Aberracla</em>), under the combination <em>Leptopsis</em> (<em>Aberracla</em>) <em>guyanensis</em> <strong>comb. nov.</strong> The SDMs for the genus <em>Leptopsis</em> showed a reasonable fit despite the genus's current limited distribution. The model indicated a wide distribution across central and northern Amazonia, with the greatest coverage in the eastern region. High-suitability areas were also identified in countries where the genus has not yet been recorded, such as Venezuela, Guyana, and Suriname. Furthermore, the model predicted high-suitability regions outside Amazonian domains, particularly in northern Colombia and Venezuela. The bioclimatic variables that contributed most to the models and probably influence the genus distribution were the precipitation of the wettest quarter and isothermality.</div></div>","PeriodicalId":49332,"journal":{"name":"Zoologischer Anzeiger","volume":"319 ","pages":"Pages 134-152"},"PeriodicalIF":1.5,"publicationDate":"2025-09-17","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"145109217","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":3,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
{"title":"The flow of non-indigenous species continues in the north Adriatic Sea: Cerastoderma edule and Mytilopsis leucophaeata (Mollusca: Bivalvia) are invading the Sacca di Goro lagoon","authors":"Giuliano Giacobelli , Valentina Tanduo , Marco Orlandi , Edoardo Turolla , Fabio Crocetta","doi":"10.1016/j.jcz.2025.09.002","DOIUrl":"10.1016/j.jcz.2025.09.002","url":null,"abstract":"<div><div>The spread of non-indigenous species (NIS) is a major threat to native biodiversity worldwide. The Mediterranean Sea is particularly affected by such a phenomenon, with high introduction rates and harmful consequences on its biota and ecosystems. Based on an integrative taxonomic approach, we hereby report the widespread occurrence of two NIS bivalves (<em>Cerastoderma edule</em> and <em>Mytilopsis leucophaeata</em>) in the Sacca di Goro Lagoon (Italy, north Adriatic Sea). The record of the former species is the first for Italy and the Mediterranean Sea <em>sensu stricto</em>, and also accounts for the first confirmed evidence of this species invading new areas worldwide, whilst that of the latter potentially clarifies the taxonomic uncertainties regarding the species that has invaded the north Adriatic until 2008. Although there are no certainties regarding their introduction pathways, <em>C. edule</em> was most likely introduced with seeding activities together with clam larvae or juveniles, while <em>M. leucophaeata</em> may have arrived through natural currents from nearby populations or amidst fouling communities of natural/anthropogenic floating objects or recreational vessels. Finally, although the local impact of these two new NIS is still unpredictable, <em>C. edule</em> can achieve extraordinarily high densities and has a structural role as a habitat engineer, influencing the distribution of surface sediments, while <em>M. leucophaeata</em> can foul and clog anthropogenic substrates and outgrow and replace benthic species. A careful monitoring of the area during the next years is therefore necessary to evaluate the consequences of these new introductions.</div></div>","PeriodicalId":49332,"journal":{"name":"Zoologischer Anzeiger","volume":"319 ","pages":"Pages 109-116"},"PeriodicalIF":1.5,"publicationDate":"2025-09-09","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"145106061","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":3,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Nanda Muhammad Razi , Sri Riska Rahayu , Luvi Syafrida Handayani , Nanda Ulfa Khaira , Muhammad Syukran , Ahmad Sazali A. Sitorus , Cut Tria Aswina , Putri Khairani Amalia Siregar , Muhammad Nasir , Nur Fadli , Martin Wilkes , Mohd Nor Siti-Azizah , Zainal Abidin Muchlisin
{"title":"DNA barcoding of fish from Singkil peat swamp waters of Aceh Province, Indonesia","authors":"Nanda Muhammad Razi , Sri Riska Rahayu , Luvi Syafrida Handayani , Nanda Ulfa Khaira , Muhammad Syukran , Ahmad Sazali A. Sitorus , Cut Tria Aswina , Putri Khairani Amalia Siregar , Muhammad Nasir , Nur Fadli , Martin Wilkes , Mohd Nor Siti-Azizah , Zainal Abidin Muchlisin","doi":"10.1016/j.jcz.2025.09.003","DOIUrl":"10.1016/j.jcz.2025.09.003","url":null,"abstract":"<div><div>Singkil peat swamp area is currently facing significant issues, including deforestation and the conversion of land into oil palm plantations. This situation threatens biodiversity and the wealth of genetic resources, potentially leading to the extinction of various terrestrial and aquatic species, including fish communities. Although there have been reports on the ichthyofauna in Singkil peat swamp, information regarding the genetic diversity of fish in this area is not available. Therefore, this study aimed to explore the use of DNA barcode method to validate the taxonomic status, as well as analyze the diversity and genetic relationships of fish found in Singkil peat swamp area, located in Aceh Province. A total of 123 DNA sequences were successfully obtained, leading to the generation of 88 haplotypes, comprising 34 species, 23 families, and 29 genera. The analysis of genetic distance was performed between various species. The results showed that the closest genetic distance of 4.04 % was between <em>Ambassis macracanthus</em> and <em>Ambassis nalua</em>, while the highest at 31.62 % was between <em>Mystus nigriceps</em> and <em>Toxotes jaculatrix</em>. <em>Cyclocheilichthys apogon</em> had the highest haplotype diversity with 6 haplotypes, while <em>Caranx tille, Oreochromis niloticus</em>, and <em>Taenioides cirratus</em> showed the lowest with 1 haplotype, respectively. Scientific data related to molecular genetics is crucial for effective fisheries resource management.</div></div>","PeriodicalId":49332,"journal":{"name":"Zoologischer Anzeiger","volume":"319 ","pages":"Pages 117-126"},"PeriodicalIF":1.5,"publicationDate":"2025-09-09","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"145106063","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":3,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
{"title":"A new species of eelpout of the genus Lycenchelys Gill from the bathyal depths of the Sea of Okhotsk","authors":"Mikhail V. Nazarkin","doi":"10.1016/j.jcz.2025.09.004","DOIUrl":"10.1016/j.jcz.2025.09.004","url":null,"abstract":"<div><div>A new species of eelpout, <em>Lycenchelys hexapora</em>, is described based on the holotype obtained from the southern part of the Sea of Okhotsk, at a depth of 1040–1050 m. The new species is characterized by a low vertebral count for its genus (89), a black stomach, and six postorbital pores, that together distinguish it from all other congeners. The new species is the sixth representative of <em>Lycenchelys</em> in the Sea of Okhotsk. The possible closest relatives of the new species are distributed hundreds kilometers away from its locality, which hints to the extensive area of their common ancestor.</div></div>","PeriodicalId":49332,"journal":{"name":"Zoologischer Anzeiger","volume":"319 ","pages":"Pages 103-108"},"PeriodicalIF":1.5,"publicationDate":"2025-09-09","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"145049753","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":3,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Jordan Hernández-Martelo , Mathias Jabs , Tamara Contador , Sanghee Kim , Sook young Lee , Laura M. Pérez , Mónica Remedios-De-León , Enrique Morelli , Peter Convey , Hugo A. Benítez
{"title":"Evolving to invade: Using geometric morphometrics to assess wing shape variation in the Antarctic non-native fly Trichocera maculipennis","authors":"Jordan Hernández-Martelo , Mathias Jabs , Tamara Contador , Sanghee Kim , Sook young Lee , Laura M. Pérez , Mónica Remedios-De-León , Enrique Morelli , Peter Convey , Hugo A. Benítez","doi":"10.1016/j.jcz.2025.09.001","DOIUrl":"10.1016/j.jcz.2025.09.001","url":null,"abstract":"<div><div>Globalization has increased human movement, transforming ecosystems and introducing non-native species that threaten biodiversity, particularly in Antarctica. Due to its harsh climatic conditions, Antarctica has largely remained protected from biological invasions. However, rising human activity and environmental changes are facilitating the introduction of non-native species. The risks posed by species like <em>Trichocera maculipennis</em> are significant, as they can adapt and survive in new environments. This highlights the importance of understanding the adaptive factors that contribute to their success in order to prevent future invasions and protect the continent's unique ecosystem. The aim of this study was to quantify the wing morphological variation of <em>T. maculipennis</em> individuals associated with different Antarctic bases, analyzing this variation within a spatial and temporal context using morphometric tools. Samples were collected during Antarctic summer campaigns (2017/18 to 2019/20 and 2022/23) at the following research stations on King George Island: Artigas (Uruguay), Bellingshausen (Russia), Profesor Julio Escudero (Chile), and King Sejong (Korea). Morphometric analyses revealed changes in wing shape among study sites, with King Sejong individuals showing the most significant differences. Additionally, a temporal shift in wing size was observed, with specimens from 2017/18 and 2018/19 having larger average wings compared to those from 2019/20 and 2021/22. These results emphasize the importance of understanding the factors contributing to the success of non-native species to prevent future invasions and protect the unique ecosystem of Antarctica.</div></div>","PeriodicalId":49332,"journal":{"name":"Zoologischer Anzeiger","volume":"319 ","pages":"Pages 70-77"},"PeriodicalIF":1.5,"publicationDate":"2025-09-03","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"144989249","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":3,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}