First records from Cape Verde and range extension of Coniophora eremophila Lindsey & Gilb. (Basidiomycota, Boletales): a morphological and molecular identification

. In the framework of a research project on corticioid fungi (Basidiomycota) from Cape Verde, we collected several specimens provisionally assigned to Coniophora sp. On the basis of morphological and molecular analyses, we identified these specimens as C. eremophila Lindsey & Gilb. These records extend the geographical distribution of this species by approximately 8,800 km in a straight line west to east, from its previously known North American locality and 8,700 km southwest to northeast from the other known locality in Chile. The pres-ence of the genus Coniophora is reported for the first time in the Cape Verde Archipelago.


Introduction
The Cape Verde archipelago is situated in the North Atlantic Ocean between 14°45′N, 022°40′W and 17° 10′N, 025°20′W and together with the Azores, Madeira, and the Canary Islands form the Macaronesian region. Cape Verde is composed of 10 volcanic islands with varying sizes, topography, and altitudes. The archipelago is spread over more than 58,000 km 2 , has a surface of 4,020 km 2 , and is located 570 km off the coast of Senegal, West Africa. The islands are usually classified in three groups: Northern Islands (Santo Antão, São Vicente, Santa Luzia, and São Nicolau), Southern Islands (Santiago, Fogo, and Brava), both characterized by hilly landscapes, and the Eastern group (Sal, Boavista, and Maio) with flat landscapes. The climate is tropical dry to semi-arid, with annual precipitations of 80-300 mm in arid coastal zones to 1,200-1,600 mm in the highlands; the mean annual temperatures range from 23-27 °C at sea level to 18-20 °C at high altitude (Duarte and Romeiras 2009).
There is little information about the mycobiota from Cape Verde. A preliminary list of non-lichenized fungi (Bañares Baudet 2005) recorded 58 species, and only seventeen basidiomycetes were included. Recently, three new species of gasteroid fungi (Crous et al. 2015(Crous et al. , 2016Martín et al. 2015) and two of corticioid fungi (Telleria et al. 2017) have been described from Santiago, Santo Antão, São Vicente, and Fogo Islands.
Coniophora DC. ex Merat is a widespread genus of corticioid fungi (Basidiomycota, Boletales, Coniophorinae; Binder and Hibbett 2006). It is characterized by the following morphology: basidioma resupinate, effused, adnate, and membranaceous; hymenophore smooth to tuberculate and margin with hyphal strands; hyphal system mono-, di-or trimitic, most generative hyphae clampless, however with scattered double or verticillate clamps especially in the wider marginal hyphae; basidia clavate, cylindrical to utriform, two or four sterigmata, without basal clamps; spores broadly ellipsoid, ellipsoid to ovoid, or subfusiform, apiculate; apical germ pore present, thick-walled, smooth, yellowish to brownish and more or less dextrinoid (Ginns 1982;Hallenberg 1985). Carlier et al. (2004) included 18 species and two varieties in the genus. Since then, one new species has been described (Boidin and Gilles 2004) and a new combination proposed (Parmasto 2005). The geographic distribution of Coniophora species is imperfectly known (Ginns 1982), and from the Macaronesian region, the genus has been reported from the Azores (Telleria et al. 2009), Madeira (Telleria et al. 2008), and the Canary Islands (Beltrán-Tejera et al. 2013; therefore, the new records reported here are the first time this genus has been found in the Cape Verde Archipelago. Coniophora eremophila was described by Lindsey & Gilbertson (1975) from the Sonoran Desert of Arizona, USA, where it was associated with brown rot on dead mesquite (Prosopis juliflora (Sw.) DC.) and other fallen debris of Olneya tesota A.Gray, Sambucus mexicana C.Presl ex DC., Chilopsis linearis (Cav.) Sweet, and Juglans major (Torr.) A.Heller. Currently this species is only known from the Sonoran Desert in North America (Lindsey and Gilbertson 1975;Gilbertson et al. 1976) and Punta Arenas, Chile, South America (Ginns 1982).

Methods
The specimens studied in this paper were collected in the Cape Verde archipelago within the framework of a project on the biogeography of corticoid fungi (CGL2009-07231). The field exploration took place on March 2010 on four islands: Santiago, Fogo, São Vicente, and Santo Antão (Figs. 1, 2). The specimens were collected by M. Dueñas (MD) and M.T. Telleria (Tell.) and deposited in the mycological collection (MA-Fungi) of the Real Jardín Botánico Herbarium, Madrid, Spain. This study included type specimens of Coniophora eremophila from Arizona (RLG 10925, holotype, BPI) and C. bimacrospora Decock, Bitew & G. Castillo from Ethiopia (MUCL 45009, holotype), and additional materials of the ARIZ and MA-Fungi herbaria ( Table 1).
The morphological analysis of the species was based on dried specimens; colors of basidioma are given according to ISCC-NBS Centroid Color Charts (Kelly and Judd 1976). Measurement was made from microscopic sections mounted in 3% KOH, Congo red, and Melzer's solutions, and examined at magnifications up to 1,250× using a differential interference contrast Olympus BX51 microscope. The length and width for 30 spores and 10 basidia were measured from each sample, and length/width ratios (Q) were calculated. Drawings were made with a Leica 2500 microscope with the help of a drawing tube. DNA extractions, ITS nrDNA amplifications, purifications, and sequencing protocols were performed as indicated by Telleria et al. (2017). Sequences obtained in this study were submitted to GenBank under the accession numbers indicated in Table 1. The newly generated ITS sequences were aligned in Se-Al v. 2.0a11 Carbon (Rambaut 2002) with homologous sequences retrieved from EMB/GenBank/DDBJ databases (Cochrane et al. 2011(Cochrane et al. , 2016 included in Table S1. The maximum-parsimony (MP) and maximum-likelihood (ML) analyses were performed as by Telleria et al. (2017) using PAUP* v. 4.0b10 for Macintosh (Swofford 2003); GTR+I+G was the model selected in PAUP to ML analysis. Two sequences of Serpula himanthioides (Fr.) P. Karst. and Serpula lacrymans (Wulfen) J. Schröt. were included as the outgroup.

Discussion
Most of the species of Coniophora cause brown rot in all kinds of wood, producing economic losses in several ways: e.g. Coniophora puteana (Schumach.) P. Karst. is an important destroyer of building timber. Matheron et al. (1992) reported Coniophora species associated with brown heartwood rot in branches and trunks on lemon  trees in Arizona, USA, that was identified a few years later as C. eremophila . This record was the first of a species of Coniophora causing serious lesions in living citrus and other fruit trees. Several studies have been carried out on the biology, control, development, characterization, and molecular biology of this disease (Bigelow et al. , 1998Gilbertson et al. 1996;Wilcox et al. 1996;Demetriou et al. 2000;Adaskaveg et al. 2001;Matheron et al. 2006). According to Mauk and Adaskaveg (2000), C. eremophila may be an opportunistic species disposed to colonize fruit trees weakened by other possible causes; our report offers additional information on this hypothesis, because it is also able to colonize other species of cultivated plants such as vines and sugarcane.
This study is first report of C. eremophila from the Cape Verde archipelago (Fig. 1) and extends the geographic distribution of this species by approximately 8,800 km in a straight line west to east from its previously known North American locality (Lindsey and Gilbertson 1975;Gilbertson et al. 1976) and 8,700 km southwest to northeast from the locality in Chile (Ginns 1982).