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Phylogeny and Evolution of the Arctium-Cousinia Complex (Compositae, Cardueae-Carduinae)
View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Digital.CSIC TAXON 58 (1) • February 2009: 153–171 López-Vinyallonga & al. • Arctium-Cousinia complex Phylogeny and evolution of the Arctium-Cousinia complex (Compositae, Cardueae-Carduinae) Sara López-Vinyallonga1,4*, Iraj Mehregan2,4, Núria Garcia-Jacas1, Olga Tscherneva3, Alfonso Susanna1 & Joachim W. Kadereit2 1 Botanical Institute of Barcelona (CSIC-ICUB), Pg. del Migdia s. n., 08038 Barcelona, Spain. *slopez@ibb. csic.es (author for correspondence) 2 Johannes Gutenberg-Universität Mainz, Institut für Spezielle Botanik und Botanischer Garten, 55099 Mainz, Germany 3 Komarov Botanical Institute, Ul. Prof. Popova 2, 197376 St. Petersburg, Russia 4 These authors contributed equally to this publication The phylogeny and evolution of the Arctium-Cousinia complex, including Arctium, Cousinia as one of the largest genera of Asteraceae, Hypacanthium and Schmalhausenia, is investigated. This group of genera has its highest diversity in the Irano-Turanian region and the mountains of Central Asia. We generated ITS and rpS4-trnT-trnL sequences for altogether 138 species, including 129 (of ca. 600) species of Cousinia. As found in previous analyses, Cousinia is not monophyletic. Instead, Cousinia subgg. Cynaroides and Hypacanthodes with together ca. 30 species are more closely related to Arctium, Hypacanthium and Schmalhausenia (Arc- tioid clade) than to subg. Cousinia (Cousinioid clade). The Arctioid and Cousiniod clades are also supported by pollen morphology and chromosome number as reported earlier. In the Arctioid clade, the distribution of morphological characters important for generic delimitation, mainly leaf shape and armature and morphology of involucral bracts, are highly incongruent with phylogenetic relationships as implied by the molecular data. -
The First Miocene Fossils of Lacerta Cf. Trilineata (Squamata, Lacertidae) with A
bioRxiv preprint doi: https://doi.org/10.1101/612572; this version posted April 17, 2019. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY 4.0 International license. The first Miocene fossils of Lacerta cf. trilineata (Squamata, Lacertidae) with a comparative study of the main cranial osteological differences in green lizards and their relatives Andrej Čerňanský1,* and Elena V. Syromyatnikova2, 3 1Department of Ecology, Laboratory of Evolutionary Biology, Faculty of Natural Sciences, Comenius University in Bratislava, Mlynská dolina, 84215, Bratislava, Slovakia 2Borissiak Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya 123, 117997 Moscow, Russia 3Zoological Institute, Russian Academy of Sciences, Universitetskaya nab., 1, St. Petersburg, 199034 Russia * Email: [email protected] Running Head: Green lizard from the Miocene of Russia Abstract We here describe the first fossil remains of a green lizardof the Lacerta group from the late Miocene (MN 13) of the Solnechnodolsk locality in southern European Russia. This region of Europe is crucial for our understanding of the paleobiogeography and evolution of these middle-sized lizards. Although this clade has a broad geographical distribution across the continent today, its presence in the fossil record has only rarely been reported. In contrast to that, the material described here is abundant, consists of a premaxilla, maxillae, frontals, bioRxiv preprint doi: https://doi.org/10.1101/612572; this version posted April 17, 2019. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. -
(Amsel, 1954) (Lepidoptera: Pyralidae, Phycitinae) – a New Species for the Croatian Pyraloid Moth Fauna, with an Updated Checklist
NAT. CROAT. VOL. 30 No 1 37–52 ZAGREB July 31, 2021 original scientific paper / izvorni znanstveni rad DOI 10.20302/NC.2021.30.4 PSOROSA MEDITERRANELLA (AMSEL, 1954) (LEPIDOPTERA: PYRALIDAE, PHYCITINAE) – A NEW SPECIES FOR THE CROATIAN PYRALOID MOTH FAUNA, WITH AN UPDATED CHECKLIST DANIJELA GUMHALTER Azuritweg 2, 70619 Stuttgart, Germany (e-mail: [email protected]) Gumhalter, D.: Psorosa mediterranella (Amsel, 1954) (Lepidoptera: Pyralidae, Phycitinae) – a new species for the Croatian pyraloid moth fauna, with an updated checklist. Nat. Croat., Vol. 30, No. 1, 37–52, 2021, Zagreb. From 2016 to 2020 numerous surveys were undertaken to improve the knowledge of the pyraloid moth fauna of Biokovo Nature Park. On August 27th, 2020 one specimen of Psorosa mediterranella (Amsel, 1954) from the family Pyralidae was collected on a small meadow (985 m a.s.l.) on Mt Biok- ovo. In this paper, the first data about the occurrence of this species in Croatia are presented. The previ- ous mention in the literature for Croatia was considered to be a misidentification of the past and has thus not been included in the checklist of Croatian pyraloid moth species. P. mediterranella was recorded for the first time in Croatia in recent investigations and, after other additions to the checklist have been counted, is the 396th species in the Croatian pyraloid moth fauna. An overview of the overall pyraloid moth fauna of Croatia is given in the updated species list. Keywords: Psorosa mediterranella, Pyraloidea, Pyralidae, fauna, Biokovo, Croatia Gumhalter, D.: Psorosa mediterranella (Amsel, 1954) (Lepidoptera: Pyralidae, Phycitinae) – nova vrsta u hrvatskoj fauni Pyraloidea, s nadopunjenim popisom vrsta. -
Two Cases of Melanism in the Ring-Headed Dwarf Snake Eirenis Modestus (Martin, 1838) from Kastellorizo, Greece (Serpentes: Colubridae)
Herpetology Notes, volume 11: 175-178 (2018) (published online on 20 February 2018) Two cases of melanism in the Ring-headed Dwarf Snake Eirenis modestus (Martin, 1838) from Kastellorizo, Greece (Serpentes: Colubridae) Konstantinos Kalaentzis1,*, Christos Kazilas1 and Ilias Strachinis1 Pigmentation serves a protective role in many 2016). A possible adaptive hypothesis for melanism in animals, including snakes, whether it functions in snakes is protection against sun damage (Lorioux et al., camouflage, warning, mimicry, or thermoregulation 2008; Jablonski and Kautman, 2017). (Bechtel, 1978; Krecsák, 2008). The observable The Ring-headed Dwarf Snake, Eirenis modestus colouration and pattern of a snake is the result of the (Martin, 1838), is a medium-sized colubrid snake presence of variously coloured pigments in specific reaching a maximum total length of 70 cm (Çiçek and places in the skin (Bechtel, 1978). Four different types Mermer, 2007). The Dwarf Snake inhabits rocky areas of pigment-bearing cells called chromatophores can with sparse vegetation and often hides under stones, be found in the skin of reptiles, namely melanophores, where it feeds mainly on terrestrial arthropods (Çiçek iridophores, erythrophores, and xanthophores (Bechtel, and Mermer, 2007). It is widely distributed (Fig. 1), 1978). Abnormalities in the pigment formation or the occurring mainly in the Caucasus (Armenia, southern interaction between the different types of pigment may Azerbaijan, eastern Georgia, southern Russia), Greece result in various chromatic disorders, which cause (on the islands of Alatonissi, Chios, Fournoi, Kalymnos, abnormal colouration of the skin and its derivatives Kastellorizo, Leros, Lesvos, Samiopoula, Samos, (Rook et al., 1998). There are many literature reports and Symi), northwestern Iran, and Turkey (Çiçek and describing chromatic anomalies in snakes, of which Mermer, 2007; Mahlow et al., 2013). -
Herpetofaunistic Diversity of the Cres-Lošinj Archipelago (Croatian Adriatic)
University of Sopron Roth Gyula Doctoral School of Forestry and Wildlife Management Sciences Ph.D. thesis Herpetofaunistic diversity of the Cres-Lošinj Archipelago (Croatian Adriatic) Tamás Tóth Sopron 2018 Roth Gyula Doctoral School of Forestry and Wildlife Management Sciences Nature Conservation Program Supervisors: Prof. Dr. Faragó Sándor Dr. Gál János Introduction In recent years the Croatian islands, especially those of the Cres-Lošinj Archipelago became the focus of research of herpetologists. However, in spite of a long interest encompassing more than a hundred years, numerous gaps remain in our herpetological knowledge. For this reason, the author wished to contribute to a better understanding by performing studies outlined below. Aims The first task was to map the distribution of amphibians and reptiles inhabiting the archipelago as data were lacking for several of the smaller islands and also the fauna of the bigger islands was insufficiently known. Subsequently, the faunistic information derived from the scientific literature and field surveys conducted by the author as well as available geological and paleogeological data were compared and analysed from a zoogeographic point of view. The author wished to identify regions of the islands boasting the greatest herpetofaunal diversity by creating dot maps based on collecting localities. To answer the question which snake species and which individuals are going to be a victim of the traffic snake roadkill and literature survey were used. The author also identified where are the areas where the most snakes are hit by a vehicle on Cres. By gathering road-killed snakes and comparing their locality data with published occurrences the author seeked to identify species most vulnerable to vehicular traffic and road sections posing the greatest threat to snakes on Cres Island. -
The Analysis of the Flora of the Po@Ega Valley and the Surrounding Mountains
View metadata, citation and similar papers at core.ac.uk brought to you by CORE NAT. CROAT. VOL. 7 No 3 227¿274 ZAGREB September 30, 1998 ISSN 1330¿0520 UDK 581.93(497.5/1–18) THE ANALYSIS OF THE FLORA OF THE PO@EGA VALLEY AND THE SURROUNDING MOUNTAINS MIRKO TOMA[EVI] Dr. Vlatka Ma~eka 9, 34000 Po`ega, Croatia Toma{evi} M.: The analysis of the flora of the Po`ega Valley and the surrounding moun- tains, Nat. Croat., Vol. 7, No. 3., 227¿274, 1998, Zagreb Researching the vascular flora of the Po`ega Valley and the surrounding mountains, alto- gether 1467 plant taxa were recorded. An analysis was made of which floral elements particular plant taxa belonged to, as well as an analysis of the life forms. In the vegetation cover of this area plants of the Eurasian floral element as well as European plants represent the major propor- tion. This shows that in the phytogeographical aspect this area belongs to the Eurosiberian- Northamerican region. According to life forms, vascular plants are distributed in the following numbers: H=650, T=355, G=148, P=209, Ch=70, Hy=33. Key words: analysis of flora, floral elements, life forms, the Po`ega Valley, Croatia Toma{evi} M.: Analiza flore Po`e{ke kotline i okolnoga gorja, Nat. Croat., Vol. 7, No. 3., 227¿274, 1998, Zagreb Istra`ivanjem vaskularne flore Po`e{ke kotline i okolnoga gorja ukupno je zabilje`eno i utvr|eno 1467 biljnih svojti. Izvr{ena je analiza pripadnosti pojedinih biljnih svojti odre|enim flornim elementima, te analiza `ivotnih oblika. -
Conserving Europe's Threatened Plants
Conserving Europe’s threatened plants Progress towards Target 8 of the Global Strategy for Plant Conservation Conserving Europe’s threatened plants Progress towards Target 8 of the Global Strategy for Plant Conservation By Suzanne Sharrock and Meirion Jones May 2009 Recommended citation: Sharrock, S. and Jones, M., 2009. Conserving Europe’s threatened plants: Progress towards Target 8 of the Global Strategy for Plant Conservation Botanic Gardens Conservation International, Richmond, UK ISBN 978-1-905164-30-1 Published by Botanic Gardens Conservation International Descanso House, 199 Kew Road, Richmond, Surrey, TW9 3BW, UK Design: John Morgan, [email protected] Acknowledgements The work of establishing a consolidated list of threatened Photo credits European plants was first initiated by Hugh Synge who developed the original database on which this report is based. All images are credited to BGCI with the exceptions of: We are most grateful to Hugh for providing this database to page 5, Nikos Krigas; page 8. Christophe Libert; page 10, BGCI and advising on further development of the list. The Pawel Kos; page 12 (upper), Nikos Krigas; page 14: James exacting task of inputting data from national Red Lists was Hitchmough; page 16 (lower), Jože Bavcon; page 17 (upper), carried out by Chris Cockel and without his dedicated work, the Nkos Krigas; page 20 (upper), Anca Sarbu; page 21, Nikos list would not have been completed. Thank you for your efforts Krigas; page 22 (upper) Simon Williams; page 22 (lower), RBG Chris. We are grateful to all the members of the European Kew; page 23 (upper), Jo Packet; page 23 (lower), Sandrine Botanic Gardens Consortium and other colleagues from Europe Godefroid; page 24 (upper) Jože Bavcon; page 24 (lower), Frank who provided essential advice, guidance and supplementary Scumacher; page 25 (upper) Michael Burkart; page 25, (lower) information on the species included in the database. -
Environmental, Socioeconomic and Cultural Heritage Baseline Page 2 of 382 Area Comp
ESIA Albania Section 6 – Environmental, Socioeconomic and Cultural Heritage Baseline Page 2 of 382 Area Comp. System Disc. Doc.- Ser. Code Code Code Code Type No. Project Title: Trans Adriatic Pipeline – TAP AAL00-ERM-641-Y-TAE-1008 ESIA Albania Section 6 - Environmental, Document Title: Rev.: 03 Socioeconomic and Cultural Heritage Baseline TABLE OF CONTENTS 6 ENVIRONMENTAL, SOCIOECONOMIC AND CULTURAL HERITAGE BASELINE 11 6.1 Introduction 11 6.2 Offshore Biological and Physical Environment 11 6.2.1 Introduction 11 6.2.2 Geographical Scope of the Baseline 13 6.2.3 Methodology and Sources of Information 13 6.2.3.1 Video Methodology 13 6.2.3.2 Environmental Survey Methodology 13 6.2.4 Legislation 15 6.2.4.1 Designated Sites 15 6.2.4.2 Sensitive and Protected Habitats / Biocenoses 16 6.2.5 Regional Overview 16 6.2.5.1 Introduction 16 6.2.5.2 Physical Environment 16 6.2.5.3 Biological Baseline 33 6.2.6 Albanian Nearshore Study Area 56 6.2.6.1 Physical Baseline 56 6.2.6.2 Biological Baseline 69 6.3 Offshore Socioeconomic Environment 73 6.3.1 Introduction 73 6.3.2 Harbours 75 6.3.2.1 Durrës Harbour 75 6.3.2.2 Vlorë Port 76 6.3.3 Marine Traffic 76 6.3.3.1 Ferry Traffic 79 6.3.4 Fishing 80 6.3.4.1 National Overview 80 6.3.5 Cultural Heritage 87 6.3.6 Marine Ammunition / Unexploded Ordnances (UXO) 88 6.4 Onshore Physical Environment 89 6.4.1 Climate and Ambient Air Quality 89 6.4.1.1 Overview 89 6.4.1.2 Climate 89 6.4.1.3 Wind 99 6.4.1.4 Ambient Air Quality 103 6.4.1.5 Key Findings and Conclusions 107 6.4.1.6 Limitations 108 6.4.2 Acoustic Environment 108 6.4.2.1 Acoustic Environment along the Pipeline Route 108 6.4.2.2 Acoustic Environment at CS03 112 6.4.2.3 Acoustic Environment at CS02 116 6.4.2.4 Limitations 120 6.4.3 Surface Water 120 6.4.3.1 Introduction 120 6.4.3.2 River Hydro-Morphology 121 6.4.3.3 Water Quality 127 6.4.3.4 Sediment Quality 137 6.4.3.5 Key Findings and Conclusions 141 Page 3 of 382 Area Comp. -
Kleiner Beitrag Zur Lepidopterenfauna Griechenlands (Insecta: Lepidoptera)
©Arbeitsgemeinschaft Österreichischer Entomologen, Wien, download unter www.biologiezentrum.at Z.Arb.Gem.Öst.Ent. 52 65-70 Wien, 30.9.2000 ISSN 0375-5223 Kleiner Beitrag zur Lepidopterenfauna Griechenlands (Insecta: Lepidoptera) Gemot EMBACHER Abstract The present paper deals with records of Lepidoptera from several districts of Greece. Records of 88 species are given. Donacaula nilotica (ZELLER, 1867), Cucullia absinthii (LINNAEUS, 1761) and Noctuajanthe (BORKHAUSEN, 1792) are recorded for the first time for the Greeke fauna. Zusammenfassung Die vorliegende Arbeit beschäftigt sich mit Nachweisen von Lepidopteren aus einigen Be- reichen Griechenlands. Es wird über 88 Arten aus verschiedenen Familien berichtet. Die Pyralide Donacaula nilotica (ZELLER, 1867) und die Noctuiden Cucullia absinthii (LINNAEUS, 1761) und Noctuajanthe (BORKHAUSEN, 1792) sind neu für die griechische Fauna. Key words: Greece, Lepidoptera, distribution, new records. Einleitung Wenn man begeisterter Griechenland-Urlauber und außerdem Lepidopterologe ist, ist es nicht verwunderlich, wenn man auch als Lokalfaunist (Fauna Salzburgs) nicht widerstehen kann, nach Möglichkeit Schmetterlinge aus seinen Urlaubsorten mit nach Hause zu neh- men. Ohne systematischen Fang zu betreiben, erhält man mit etwas Glück doch auch gele- gentlich Tiere, die eventuell für Spezialisten und Faunenbearbeiter interessant oder für ge- wisse geographische Bereiche sogar Neufunde sind. Die folgende Liste soll dazu ein klei- ner Beitrag sein. Artenliste Zygaenidae Jordanita notata (ZELLER, 1847): Kreta, Stalis, 13.4.1990, ein è morgens an einer Straßen- lampe. Adscita mannii (LEDERER, 1853): Thassos, Chrissi Amoudia, 4.- 10.6.1999, einzeln. Zygaena loti ([DENIS & SCHIFFERMÜLLER], 1775): Thassos, Chrissi Amoudia, 4.-10.6.1999, zahlreich. 65 ©Arbeitsgemeinschaft Österreichischer Entomologen, Wien, download unter www.biologiezentrum.at Zygaena fìlipendulae (LINNAEUS, 1758): Zakynthos, Argassi, 25.5.1998. -
Hortus Botanicus Universitatis Posnaniensis Index Seminum
HORTUS BOTANICUS UNIVERSITATIS POSNANIENSIS INDEX SEMINUM 2020-2021 ANNO 2020 COLLECTORUM QUAE HORTUS BOTANICUS UNIVERSITATIS POSNANIENSIS MUTUO COMMUTANDA OFFERT OGRÓD BOTANICZNY UNIWERSYTETU IM. ADAMA MICKIEWICZA UL. DĄBROWSKIEGO 165 PL – 60-594 POZNAŃ ebgconsortiumindexseminum2020 ebgconsortiumindexseminum2021 Information Informacja Year of foundation – 1925 Rok założenia – 1925 Area about 22 ha, including about 800 m2 of greenhouses Aktualna powierzchnia około 22 ha w tym około 800 m2 pod szkłem Number of taxa – about 7500 Liczba taksonów – około 7500 1. Location: 1. Położenie: the Botanical Garden of the A. Mickiewicz University is situated in the W part of Poznań zachodnia część miasta Poznania latitude – 52o 25‘N szerokość geograficzna – 52o 25‘N longitude – 16o 55‘E długość geograficzna – 16o 55‘E the altitude is 89.2 m a.s.l. wysokość n.p.m. – 89.2 m 2. The types of soils: 2. Typy gleb: – brown soil – brunatna – rot soil on mineral ground – murszowa na podłożu mineralnym – gray forest soil – szara gleba leśna SEMINA PLANTARUM EX LOCIS NATURALIBUS COLLECTA zbierał/collected gatunek/species stanowisko/location by MAGNOLIOPHYTA Magnoliopsida Apiaceae 1. Daucus carota L. PL, prov. Wielkopolskie, Poznań, Szczepankowo J. Jaskulska 2. Peucedanum oreoselinum (L.) Moench PL, prov. Kujawsko-Pomorskie, Folusz J. Jaskulska Asteraceae 3. Achillea millefolium L. s.str. PL, prov. Wielkopolskie, Kamionki J. Jaskulska 4. Achillea millefolium L. s.str. PL, prov. Wielkopolskie, Koninko J. Jaskulska 5. Artemisia vulgaris L. PL, prov. Wielkopolskie, Kamionki J. Jaskulska 6. Artemisia vulgaris L. PL, prov. Wielkopolskie, Koninko J. Jaskulska 7. Bidens tripartita L. PL, prov. Wielkopolskie, Koninko J. Jaskulska 8. Centaurea scabiosa L. PL, prov. Kujawsko-Pomorskie, Folusz J. -
Environmental Impacts
E4482 V4 Public Enterprise for State Roads Environmental and Social Assessment Report Public Disclosure Authorized PUBLIC ENTERPRISE FOR STATE ROADS Republic of Macedonia Public Disclosure Authorized NATIONAL AND REGIONAL ROADS REHABILITATION PROJECT Final Environmental and Social Assessment Report (ESAR) and Environmental Management Plan (EMP) for rehabilitation of State road R1202, Section Boshkov Most - Debar Public Disclosure Authorized Public Disclosure Authorized March 2014 Geing Krebs und Kifer International and others ltd 1 Public Enterprise for State Roads Environmental and Social Assessment Report Table of contents 1. EXECUTIVE SUMMARY AND CONCLUSION ............................... 3 2. POLICY, LEGAL AND ADMINISTRATIVE FRAMEWORK ........................ 4 3. PROJECT DESCRIPTION ........................................... 7 4. BASELINE DATA ................................................ 9 LOCATION ST.NIKOLA - LOCATION IN MAVROVO WHERE NECROPOLIS FROM THE LATE MIDDLE AGE WAS DISCOVERED. THIS LOCATION IS NEAR THE VILLAGE CHURCH ST.NIKOLA, AND UNDER THE ROAD THERE ARE STONE PLATES FROM GRAVES AND ARCHAIC CROSSES, MADE FROM LIMESTONE. ........................................................................................................................................................ 19 5. ENVIRONMENTAL IMPACTS ........................................ 20 5.1 AIR EMISSIONS ................................................................................................................................................... 20 5.2 WATER -
Nuclear and Plastid DNA Phylogeny of the Tribe Cardueae (Compositae
1 Nuclear and plastid DNA phylogeny of the tribe Cardueae 2 (Compositae) with Hyb-Seq data: A new subtribal classification and a 3 temporal framework for the origin of the tribe and the subtribes 4 5 Sonia Herrando-Morairaa,*, Juan Antonio Callejab, Mercè Galbany-Casalsb, Núria Garcia-Jacasa, Jian- 6 Quan Liuc, Javier López-Alvaradob, Jordi López-Pujola, Jennifer R. Mandeld, Noemí Montes-Morenoa, 7 Cristina Roquetb,e, Llorenç Sáezb, Alexander Sennikovf, Alfonso Susannaa, Roser Vilatersanaa 8 9 a Botanic Institute of Barcelona (IBB, CSIC-ICUB), Pg. del Migdia, s.n., 08038 Barcelona, Spain 10 b Systematics and Evolution of Vascular Plants (UAB) – Associated Unit to CSIC, Departament de 11 Biologia Animal, Biologia Vegetal i Ecologia, Facultat de Biociències, Universitat Autònoma de 12 Barcelona, ES-08193 Bellaterra, Spain 13 c Key Laboratory for Bio-Resources and Eco-Environment, College of Life Sciences, Sichuan University, 14 Chengdu, China 15 d Department of Biological Sciences, University of Memphis, Memphis, TN 38152, USA 16 e Univ. Grenoble Alpes, Univ. Savoie Mont Blanc, CNRS, LECA (Laboratoire d’Ecologie Alpine), FR- 17 38000 Grenoble, France 18 f Botanical Museum, Finnish Museum of Natural History, PO Box 7, FI-00014 University of Helsinki, 19 Finland; and Herbarium, Komarov Botanical Institute of Russian Academy of Sciences, Prof. Popov str. 20 2, 197376 St. Petersburg, Russia 21 22 *Corresponding author at: Botanic Institute of Barcelona (IBB, CSIC-ICUB), Pg. del Migdia, s. n., ES- 23 08038 Barcelona, Spain. E-mail address: [email protected] (S. Herrando-Moraira). 24 25 Abstract 26 Classification of the tribe Cardueae in natural subtribes has always been a challenge due to the lack of 27 support of some critical branches in previous phylogenies based on traditional Sanger markers.