<?xml version="1.0" encoding="UTF-8"?><!DOCTYPE article  PUBLIC "-//NLM//DTD Journal Publishing DTD v3.0 20080202//EN" "http://dtd.nlm.nih.gov/publishing/3.0/journalpublishing3.dtd"><article xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" dtd-version="3.0" xml:lang="en" article-type="research article"><front><journal-meta><journal-id journal-id-type="publisher-id">AJPS</journal-id><journal-title-group><journal-title>American Journal of Plant Sciences</journal-title></journal-title-group><issn pub-type="epub">2158-2742</issn><publisher><publisher-name>Scientific Research Publishing</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="doi">10.4236/ajps.2017.84062</article-id><article-id pub-id-type="publisher-id">AJPS-75089</article-id><article-categories><subj-group subj-group-type="heading"><subject>Articles</subject></subj-group><subj-group subj-group-type="Discipline-v2"><subject>Biomedical&amp;Life Sciences</subject></subj-group></article-categories><title-group><article-title>
 
 
  Novelties in &lt;i&gt;Campyloneurum&lt;/i&gt; (Polypodiaceae) from Mesoamerica
 
</article-title></title-group><contrib-group><contrib contrib-type="author" xlink:type="simple"><name name-style="western"><surname>Alexander</surname><given-names>Francisco Rojas-Alvarado</given-names></name><xref ref-type="aff" rid="aff1"><sub>1</sub></xref></contrib></contrib-group><aff id="aff1"><label>1</label><addr-line>Universidad Nacional de Costa Rica, Heredia, Costa Rica</addr-line></aff><author-notes><corresp id="cor1">* E-mail:</corresp></author-notes><pub-date pub-type="epub"><day>03</day><month>03</month><year>2017</year></pub-date><volume>08</volume><issue>04</issue><fpage>921</fpage><lpage>940</lpage><history><date date-type="received"><day>February</day>	<month>27,</month>	<year>2017</year></date><date date-type="rev-recd"><day>Accepted:</day>	<month>March</month>	<year>28,</year>	</date><date date-type="accepted"><day>March</day>	<month>31,</month>	<year>2017</year></date></history><permissions><copyright-statement>&#169; Copyright  2014 by authors and Scientific Research Publishing Inc. </copyright-statement><copyright-year>2014</copyright-year><license><license-p>This work is licensed under the Creative Commons Attribution International License (CC BY). http://creativecommons.org/licenses/by/4.0/</license-p></license></permissions><abstract><p>
 
 
  Three new species are described and another is recognized in 
  Campyloneurum
  . Two species occur in Mexico to El Salvador and the others two are present in Nicaragua, Costa Rica and Panama. The new species are 
  Campyloneurum acutum 
  A. Rojas,
   C. leoniae
   A. Rojas and 
  C. madrense
   A. Rojas. The first species differs from 
  C. xalapense
   by broader rhizome scales, acute to attenuate blade apex, commonly two series of areoles between parallel veins, commonly one sori per areole and hydathodes absent or few evident. 
  Campyloneurum leoniae
   differs from 
  C. irregulare
   by more compact and non
  -
  pruinose rhizomes, thinner stipes, relatively broader, thinner and more opaque laminae, and distribution at lower elevations. Campyloneurum madrense is different to C. angustifolium because 
  it 
  has narrower and darker rhizome scales with denticulate margin, long scales cells and distributed at lower elevations. Finally, Campylonerum pittieri is here recognized as valid species and it differs from the similar C. amphostenon by longer rhizomes with appressed scales that have a patent, long acuminate apices, relative broader laminae, veins evident and distribution at higher elevations.
 
</p></abstract><kwd-group><kwd>&lt;i&gt;Campyloneurum amphostenon&lt;/i&gt; Complex</kwd><kwd> &lt;i&gt;Campyloneurum angustifolium&lt;/i&gt; Complex</kwd><kwd> &lt;i&gt;Campyloneurum xalapense&lt;/i&gt;</kwd><kwd> Lectotype Designed</kwd><kwd> New Species</kwd><kwd> Ferns</kwd><kwd> Species Recognized</kwd></kwd-group></article-meta></front><body><sec id="s1"><title>1. Introduction</title><p>Campyloneurum C. Presl is a Neotropical genus with about 50 species [<xref ref-type="bibr" rid="scirp.75089-ref1">1</xref>] . This genus is recognized by having sterile and fertile monomorphic leaves, single, entire and glabrous blades or with few scales on the rachis, sori in 2 - 4 rows between the main lateral veins and they born on the tips of included veinlets [<xref ref-type="bibr" rid="scirp.75089-ref2">2</xref>] .</p><p>In the Campyloneurum angustifiolium (Sw.) F&#233;e complex, Lellinger described C. centrobrasilianum Lellinger, C. cooperi Lellinger and C. irregulare Lellinger. The first species was compared with C. angustipaleatum (Alston) M. Mey. ex Lellinger, the second one with C. rigidum J. Sm. and the third with C. amphostenon (Kunze ex Klotzsch) F&#233;e. Also he made a new combination from Polypodium angustifolium f. densifolium Hieron. as C. densifolium (Hieron.) Lellinger [<xref ref-type="bibr" rid="scirp.75089-ref3">3</xref>] . Of the species previously mentioned, Lellinger registered from Costa Rica to Campyloneurum cooperi and C. irregulare [<xref ref-type="bibr" rid="scirp.75089-ref3">3</xref>] . However, Le&#243;n synonymized this species joining to C. angustifolium s. l. and mentioned that the group require more studies. Additionally, Le&#243;n registered to C. angustipaleatum (Alston) M. Mey. ex Lellinger and C. densifolium from Costa Rica [<xref ref-type="bibr" rid="scirp.75089-ref4">4</xref>] .</p><p>Lellinger mentioned two distributional ranges from Campyloneurum irregulare in Costa Rica, between 800 - 1500 m and between 2200 - 3300 m [<xref ref-type="bibr" rid="scirp.75089-ref5">5</xref>] . Rojas found that really are two entities with different geographic and altitudinal distribution. Since the plants with the northernmost distribution between Costa Rica and Nicaragua present a non-farinosous rhizome, rhizome scales are with basal cells less markedly irregular and distribute to (700-) 1200 - 1700 m, while the typical form with pruinose rhizome and basal cells of the rhizome scales markedly irregular is located to (1200-) 1700 - 2500 m. Rojas also mentioned that lowland plants need to merit consideration of variety or species [<xref ref-type="bibr" rid="scirp.75089-ref6">6</xref>] .</p><p>Mickel &amp; Smith made difference between C. angustifolium and C. ensifolium (Willd.) J. Sm., principally referring to rhizome scales, the first with long acuminate apex and with oblong cells, the other with rounded to short acute apex and the cells rounded to squarish [<xref ref-type="bibr" rid="scirp.75089-ref7">7</xref>] .</p><p>Respect to Campyloneurum xalapense F&#233;e, Le&#243;n mentioned that this taxon has differences in the blade form and variation in rhizome scales between lanceolate to linear, related with populations but not between characters [<xref ref-type="bibr" rid="scirp.75089-ref8">8</xref>] . Mickel &amp; Smith mentioned different chromosome number between Oaxaca (=74) and Costa Rica (=148) [<xref ref-type="bibr" rid="scirp.75089-ref7">7</xref>] , probably due because they are different taxa.</p><p>Le&#243;n defined to Campyloneurum amphostenon (Kunze ex Klotzsch) F&#233;e as plants with long-creeping rhizome with dark brown and ovate scales, they are appressed at base, with patent apex [<xref ref-type="bibr" rid="scirp.75089-ref4">4</xref>] [<xref ref-type="bibr" rid="scirp.75089-ref8">8</xref>] . Respect to same species, Rojas found two different entities and separated them in his <xref ref-type="table" rid="table2">Table 2</xref>, [<xref ref-type="bibr" rid="scirp.75089-ref6">6</xref>] , relating entity 1 to the entity proposed by Le&#243;n [<xref ref-type="bibr" rid="scirp.75089-ref4">4</xref>] [<xref ref-type="bibr" rid="scirp.75089-ref8">8</xref>] and entity 2 as a possible hybrid that has short-creeping rhizome with scales 1.5 - 3 times longer than wide and brown- yellowish, fronds relatively shorter and narrower, veins few evident and distributed at low elevation.</p><p>Le&#243;n [<xref ref-type="bibr" rid="scirp.75089-ref8">8</xref>] mentioned 18 species of Campyloneurum from Mesoamerica, but Rojas with the publication of C. gracile and the recognition of C. cooperi and C. irregulare [<xref ref-type="bibr" rid="scirp.75089-ref6">6</xref>] , the inclusion of Hyalotrichopteris anetioides (Christ) W.H. Wagner in Campyloneurum [<xref ref-type="bibr" rid="scirp.75089-ref9">9</xref>] and four species that are included in this work; the current number of species for the region arrives to 26.</p></sec><sec id="s2"><title>2. Materials and Methods</title><p>The new species here considered are the result of comparisons with specimens of other Mexican and Central American species, and a review of related species and keys from the neotropical ferns by G&#243;mez &amp; Arbel&#225;ez [<xref ref-type="bibr" rid="scirp.75089-ref1">1</xref>] , Lellinger [<xref ref-type="bibr" rid="scirp.75089-ref5">5</xref>] , Mickel &amp; Smith [<xref ref-type="bibr" rid="scirp.75089-ref7">7</xref>] , Mickel &amp; Beitel [<xref ref-type="bibr" rid="scirp.75089-ref10">10</xref>] , Smith [<xref ref-type="bibr" rid="scirp.75089-ref11">11</xref>] and Stolze [<xref ref-type="bibr" rid="scirp.75089-ref12">12</xref>] and other papers in the Campyloneurum genus [<xref ref-type="bibr" rid="scirp.75089-ref3">3</xref>] [<xref ref-type="bibr" rid="scirp.75089-ref6">6</xref>] . The examined specimens are deposited in the following herbaria CHAPA, CR, MEXU, MO, UC and XAL [<xref ref-type="bibr" rid="scirp.75089-ref13">13</xref>] . To ensure the correct application names, original type material or digital type images were examined as available (Jstor Global Plants (http://plants.jstor.org/)), and the new names were corroborated with International Plant Name Index (http://www.ipni.org/ipni/plantnamesearchpage.do).</p></sec><sec id="s3"><title>3. Results</title><sec id="s3_1"><title>3.1. New Species</title><sec id="s3_1_1"><title>3.1.1. Campyloneurum acutum A. Rojas, Sp. Nov. (<xref ref-type="fig" rid="fig1">Figure 1</xref>)</title><p>TYPE: MEXICO. Guerrero: Mpio. Atoyac, a 8 km al SW de Puerto del Gallo, camino a Atoyac, 2140 m, 16 jun 1982, E. Mart&#237;nez 982 (holotype: MEXU).</p><p>Diagnoses: Campyloneurum acutum differs from C. xalapense by lanceolate and broader rhizome scales, acute blade apex and parallel veins with commonly two series of areoles.</p><p>Description: Epiphytic; rhizome 3 - 6 mm in diameter, short-creeping, fronds 2 - 5 mm distant between them, not pruinose, brown to blackish when dry; rhizome scales 3 - 6 &#215; 0.5 - 1.5 mm, lanceolate, brown to yellowish-brown, clathrate or scarcely so with transparent lumina, marginally entire and acute at tip; fronds (20-) 45 - 93 cm long, moderately decumbent; stipe (3-) 5 - 9 cm long, 1/12 - 1/9 of the frond length, strawish to pale brown, glabrous, phyllopodia 2 - 3 mm long; blade (17-) 40 - 84 &#215; (2-) 3 - 4.3 (-5.5) cm, narrowly elliptic to linear-oblong, gradually reduced at base, long acute to attenuate at apex, chartaceous, lustrous; hydathodes absent or small; rachis strawish to pale brown, glabrous; veins few evident, commonly forming 2 series of areoles between parallel veins, included veinlets one (rarely 2) per areole; sori rounded, in 4 - 6 rows between costa and margin.</p><p>Etimology: The name of the new species refers to the acute blade apex.</p><p>Distribution: Known from Mexico and El Salvador at 1000 - 2700 m elevation.</p><p>Additional revised specimens (paratypes): MEXICO. Chiapas: Municipio San Andr&#233;s Larrainzar, near the summit of Chuchil Ton, NE of Bochil, 2700 m, 3 Aug 1972, D. Breedlove 26780-A (MEXU); Municipio Motozintla de Mendoza, 45 - 50 km NE of Huixtla along road to Motozintla, 1900 m, 17 Nov 1971, D. Breedlove &amp; A. Smith 22658 (MEXU); SE side of Volc&#225;n Tacan&#225; above Talquian, 2200 m, 16 Jan 1973, D. Breedlove &amp; A. Smith 31631 (MEXU); to 22 km E of Lagunas de Monte Bello, over the trail to Santa Elena, 22 Nov 1982, E. Cabrera &amp; H. de Cabrera 3687 (MEXU); above El Rosario, 8 mi S of Motozintla, 1800 m, 10 Jul 1977, T. Croat 40731 (MEXU, MO), ib&#237;dem, T. Croat 40732 (MEXU, MO); Municipio Jaltenango, El Triunfo Reserve, trail NNW from El Triunfo camp towards Palo Gordo camp, 1 - 3 km from El Triunfo camp, 15˚39'N, 92˚50'W, 2000 m, 21 Feb 1990, R. Hampshire et al. 530 (MEXU); Municipio Jaltenango, Reserva El Triunfo, 1 polygon, HQ camp, 15˚39'N, 92˚48'W, 1850 m, 13 Jun 1990, M. Heath &amp; A. Long 976 (MEXU); Municipio La Concordia, Cerro Ceb&#250;, polygon, Core zone 5, 15˚49'24.4''N, 93˚02'44''W, 2200 m, 17 jul 2005, J. L&#243;pez 49 (MEXU); Municipio Mapastepec, Triunfo-Tomatal camp, 15˚38'N, 92˚50'W, 1728 m, M. L&#243;pez 85 (MEXU); Municipio Uni&#243;n de Ju&#225;rez, between Tonin&#225; and Talqui&#225;n, 1700 - 2700 m, 10 May 1987, E. Mart&#237;nez 20884 (MEXU); Municipio Ziltepec, Pico de Loro range, 8 km S of Nueva Argentina, 15˚28'50''N, 92˚36'20''W, 2000 m, 27 Jan 1999, M. P&#233;rez 1739 (MEXU); Municipio Monte Cristo, trail to San Luis, 2 km from Laguna El Cofre, buffer zone, 15˚37'33.8''N, 92˚37'45.1''W, 2010 m, 15 Jun 2005, M. Reynoso 209 (MEXU); Municipio Tapachula, El Dormitorio, hillside W from Volc&#225;n Tacan&#225;, 22 May 1984, P. Tenorio et al. 5913 (MO). Guerrero: Municipio Atoyac, 8 km SW from Puerto del Gallo, trail to Atoyac, 2140 m, 16 Jun 1982, E. Mart&#237;nez 980A (MEXU); Reserva El Triunfo, 1900 m, 1 Jun 1987, E. Mart&#237;nez et al. 21547 (MEXU); Municipio La Independencia, Colonia Francisco Madero, near the road, 14 Mar 1981, M. Sousa et al. 1935 (MEXU); Municipio Yajal&#243;n, Surroundings of Ejido L&#225;zaro C&#225;rdenas, 17.21463˚N, 92.6059˚W, 1152 m, 11 abr 2010, G. Salazar et al. 8904 (MEXU); in Las Golondrinas, 29 km NE from Para&#237;so, trail to Puerto del Gallo, 1810 m, 5 Jun 1983, J. Soto et al. 5188 (MEXU); Municipio Zihuatanejo, 86 km NE from Zihuatanejo, by road Zihuatanejo-Ciudad Altamirano, 1530 m, 21 May 1982, P. Tenorio et al. 422 (MEXU). Oaxaca: Municipio Santiago Comaltepec, 2.8 km in a straight line to S (225˚) from La Esperanza, 17˚36'32''N, 96˚24'14''W, 1600 m, 10 Jan 1995, C. Gallardo et al. 1323 (MEXU); Municipio San Felipe Usila, on the periphery of the community of Santa Cruz Tepetotutla, 1000 m, 07 May 1995, M. Romero 2926 (MEXU); Distrito Ixtl&#225;n, Municipio Comaltepec, Vista Hermosa, 26.6 km SW from Valle Nacional, road Tuxtepec-Oaxaca, 17˚43'N, 96˚20'W, 1460 m, 26 Jan 1988, R. Torres &amp; E. Mart&#237;nez 11417 (MEXU). Queretaro: Municipio Jalpan, 2 - 3 km N from La Parada, 1250 m, 08 Mar 1991, B. Serv&#237;n 868 (MEXU). Tabasco: Municipio Tacotalpa, Sun&#250; and Patastal, 16 Feb 2000, G. Gaspar &amp; G. Oca&#241;a 94 (MEXU). Veracruz: Municipio San Andr&#233;s Tuxtla, Ejido Barrio Lerdo, slopes of Volc&#225;n San Mart&#237;n Tuxtla, 18˚34'N, 95˚10'W, 1000 m, 11 Aug 2005, T. Kr&#246;mer &amp; A. Acebey 2474 (MEXU); Municipio San Andr&#233;s Tuxtla, Ejido Ruiz Cortines, sidewalk near the town that connects with dirt road, 18˚31'59''N, 95˚08'59''W, 1100 m, 25 May 2001, A. Rinc&#243;n et al. 2427 (MEXU); Municipio Chocaman, Tepejilotla, 1300 m, 29 Jun 1979, F. Ventura 16285 (MEXU).</p><p>EL SALVADOR. Santa Ana: Cerro Montecristo, ca. 14 mi NE of Metap&#225;n, 1950 - 2000 m, 31 Jul 1977, T. Croat 42399 (MEXU, MO).</p><p>Campyloneurum acutum differs from C. xalapense by lanceolate (vs. linear or at less with long and narrow apex) and broader apex (0.5 - 1 mm vs. 0.2 - 0.5 mm) rhizome scales, acute to attenuate (vs. long acuminate to cuspidate) blade apex, hydathodes absent or small (vs. markerly evident), commonly two (vs. one) series of areoles between parallel veins and commonly one sori per areola (vs. 2). Also the new species is related to C. tenuipes but differs of it by longer (1/12 - 1/9 of the frond length vs. ca. 1/3) stipe, narrower ((2-) 3 - 4.3 (-5.5) vs. 4 - 9 cm) blade, acute to attenuate (vs. long acuminate to cuspidate) blade apex and 4 - 6 (vs. 6 - 10) sori rows between costa and margin (see <xref ref-type="fig" rid="fig1">Figure 1</xref> and <xref ref-type="table" rid="table1">Table 1</xref>).</p></sec><sec id="s3_1_2"><title>3.1.2. Campyloneurum leoniae A. Rojas, Sp. Nov. (<xref ref-type="fig" rid="fig2">Figure 2</xref>)</title><p>TYPE: COSTA RICA. Guanacaste: Cordillera de Tilar&#225;n, 1 km N of Las Nubes de R&#237;o Chiquito, Zona Monteverde, Atlantic slope, 10˚22'N, 84˚51'W, 1300 m, 4 Sep 1988, W. Haber &amp; W. Zuchowski 8691 (holotype: CR!; isotype: MO?).</p><p>Diagnoses: Campyloneurum leoniae differs from C. irregulare by more compact and non pruinose (vs. pruinose) rhizomes, relatively broader, thinner and more opaque (vs. lustrous) laminae, and distribution at lower elevations.</p><fig id="fig1"  position="float"><label><xref ref-type="fig" rid="fig1">Figure 1</xref></label><caption><title> (a)-(b). Type material of Campyloneurum acutum A. Rojas (E. Mart&#237;nez 982, MEXU). (a) Habit; (b) Rhizome scales; (c) Blade detail</title></caption><graphic mimetype="image"   position="float"  xlink:type="simple"  xlink:href="http://html.scirp.org/file/19-2603096x2.png"/></fig><table-wrap id="table1" ><label><xref ref-type="table" rid="table1">Table 1</xref></label><caption><title> Comparison of morphological characteristics among three species of Campyloneurum: C. acutum, C. tenuipes and C. xalapense</title></caption><table><tbody><thead><tr><th align="center" valign="middle" >Character</th><th align="center" valign="middle" >C. acutum</th><th align="center" valign="middle" >C. tenuipes</th><th align="center" valign="middle" >C. xalapense</th></tr></thead><tr><td align="center" valign="middle" >Long of rhizome scales</td><td align="center" valign="middle" >3 - 6 mm</td><td align="center" valign="middle" >5 - 8 mm</td><td align="center" valign="middle" >3 - 5 mm</td></tr><tr><td align="center" valign="middle" >Broad of rhizome scales</td><td align="center" valign="middle" >0.5 - 1.5</td><td align="center" valign="middle" >1 - 1.5</td><td align="center" valign="middle" >0.2 - 0.5</td></tr><tr><td align="center" valign="middle" >Color of rhizome scales</td><td align="center" valign="middle" >yellowish-brown to brown</td><td align="center" valign="middle" >brown</td><td align="center" valign="middle" >dark brown to blackish</td></tr><tr><td align="center" valign="middle" >Lumen color of rhizome scales</td><td align="center" valign="middle" >translucid</td><td align="center" valign="middle" >translucid</td><td align="center" valign="middle" >brown</td></tr><tr><td align="center" valign="middle" >Size of fronds</td><td align="center" valign="middle" >(20-) 45 - 93 cm</td><td align="center" valign="middle" >30 - 80 cm</td><td align="center" valign="middle" >(20-) 30 - 60 cm</td></tr><tr><td align="center" valign="middle" >Long of stipe</td><td align="center" valign="middle" >1/12 - 1/9 of the frond length</td><td align="center" valign="middle" >1/3 of the frond length</td><td align="center" valign="middle" >1/10 of the frond length</td></tr><tr><td align="center" valign="middle" >Broad of blade</td><td align="center" valign="middle" >(2-) 3 - 4.3 (-5.5) cm</td><td align="center" valign="middle" >4 - 9 cm</td><td align="center" valign="middle" >3 - 5.5 cm</td></tr><tr><td align="center" valign="middle" >Blade apex</td><td align="center" valign="middle" >acute to attenuate</td><td align="center" valign="middle" >acuminate to cuspidate</td><td align="center" valign="middle" >acuminate to cuspidate</td></tr><tr><td align="center" valign="middle" >Rows of areoles between parallel veins</td><td align="center" valign="middle" >commonly two</td><td align="center" valign="middle" >two</td><td align="center" valign="middle" >one</td></tr><tr><td align="center" valign="middle" >Rows of sori between midvein and margin</td><td align="center" valign="middle" >4 - 6</td><td align="center" valign="middle" >6 - 10</td><td align="center" valign="middle" >4 - 7</td></tr><tr><td align="center" valign="middle" >Altitudinal distribution</td><td align="center" valign="middle" >1000 - 2700 m</td><td align="center" valign="middle" >1000 - 2300 m</td><td align="center" valign="middle" >500 - 1700 (-2300) m</td></tr></tbody></table></table-wrap><p>Description: Epiphytic; rhizome 3 - 5 mm in diameter, short-creeping, fronds approximate to distant to 3 mm, not pruinose, blackish and glabrescent with age; rhizome scales 1.5 - 3.5 (-4) &#215; 0.8 - 1.5 mm, ovate-lanceolate to lanceolate, dark brown, clathrate, dense, cordiform at base, cells with irregular base, with the cells at scale apex 1.5 - 3 times longer than wide, lumen pale brown; fronds (8-) 15 - 35 (-45) cm long; stipe 0.5 - 4 (-5) cm long, 1/12 - 1/8 of the frond length, 1 - 1.5 mm wide (to 2 mm with wing), slight to strongly flattened, strawish to pale brown, winged throughout, glabrous or sparsely scaly, the scales 0.5 - 1.5 x ca. 0.5 mm, ovate, brown; phyllopodia 0.5 - 1 (-2) mm long; blade (7-) 13 - 30 (-40) &#215; (0.5-) 1 - 2 (-2.3) cm, linear-elliptic, attenuate on both sides, coriaceous, opaque, with commonly revolute margin; rachis strawish to brown, glabrous or sparsely scaly at base; lateral veins few evident; sori rounded, in 1 - 3 (-4) irregular rows between costa and margin.</p><p>Etimology: This new species is dedicated to Blanca Le&#243;n, Peruvian pteridologist, who has made several contributions to Campyloneurum studies.</p><p>Distribution: Known from Nicaragua and Costa Rica at 700 - 1800 (-2000) m elevation.</p><p>Additional revised specimens (paratypes): NICARAGUA. Estel&#237;: Cerro Quiab&#250;, NW of Estel&#237;, 13˚07'N, 86˚26'W, 1600 m, 24 Jul 1980, P. Moreno 1330 (CR, HNMN); foothill of Quiab&#250;, side N, 13˚06'N, 86˚25'W, 1460 - 1480 m, 2 Jul 1982, P. Moreno 16775 (CR, HNMN). Granada: Volc&#225;n Mombacho, 1345 m, 31 Jan 1980, M. Araquistain &amp; P. Moreno 1206 (CR, HNMN). Jinotega: Santa Gertrudis, 30 km E of Jinotega, 1000 m, 20 Jan 1980, M. Araquistain &amp; P. Moreno 867 (CR, HNMN); ca. 23.5 km (by road) NE of hwy. 1 at Estel&#237; and ca. 2.8 km SE of Laguna Miraflores, ca. 13˚14'N, 86˚14'W, 1400 m, 11 Jun 1981, J. Henrich &amp; W. Stevens 408 (CR, MO); ib&#237;dem, J. Henrich &amp; W. Stevens 410 (CR, MO); along hwy 3, ca. 1.9 km NW of Aranjuez, road to entrance, ca. 13&#186;02ʹN, 85˚56'W, 1460 - 1480 m, 24 Dec 1977, W. Stevens &amp; B. Krukoff 5537 (CR, MO); ca. 0,6 km from hwy 3 on road to Aranjuez, ca. 13˚01'N, 85˚55'W, ca. 1430 m, 9 Apr 1978, W. Stevens &amp; B. Krukoff 7543 (CR, MO); along hwy 3, ca 1 km NW of La Fundadora entrance, unnamed peak ca. 500 m W of hwy, ca. 13˚01'N, 85˚56'W, 1450 - 1520 m, 24 May 1981, W. Stevens et al. 20391 (CR); Miraflores, Aug 1975, J. Villa s.n. (CR). Madriz: Cerro Pataste, ca. 20 km SW of Somoto city, ca. 1700 m, 28 Aug 1982, A. Grijalva 897 (CR, HNMN). Matagalpa: La Carlota, road Matagalpa-Tuma, 1400 - 1500 m, Oct 1975, L. G&#243;mez et al. 6298 (CR); Fuente Pura, Santa Mar&#237;a de Ostuma, between Matagalpa and Jinotega, 1400 m, 26 Aug 1982, E. Mart&#237;nez et al. 1731 (MEXU); trail to Cerro La Carlota, 2 km of the trail to Tuma, 12˚58'N, 85˚52'W, 1040 - 1100 m, 5 Mar 1982, P. Moreno 15618 (CR, HNMN); Santa Mar&#237;a de Ostuma estate, 10 km N of Matagalpa, 1300 m, 17 Jul 1978, P. Vincelli 755 (CR).</p><p>COSTA RICA. Alajuela: Piedades Norte (Los &#193;ngeles) of San Ram&#243;n, 6 Mar 1929, A. Brenes 6724 (CR); La Palma of San Ram&#243;n, 2 Nov 1929, A. Brenes 7396 (145) (CR); San Pedro of San Ram&#243;n, 31 Jan 1936, A. Brenes 21558 (8) (CR); San Carlos, 14 km NE of Ciudad Quesada, 1250 m, 22 Jul 1963, A. Jim&#233;nez 950 (CR); above Angel Falls, R&#237;o La Paz Grande, on road to Sarapiqu&#237;, 1460 m, 23 Mar 1969, R. Lent 1501 (CR); above R&#237;o Toro, 10˚12'N, 84˚19'W, 1550 m, 3 Sep 1972, R. Lent 2812 (CR); La Palma of San Ram&#243;n, 3 - 10 Aug 1935, M. Quir&#243;s 259 (CR); Carrillos de Po&#225;s, 23 Aug 1936, M. Quir&#243;s 569 (CR); San Ram&#243;n, Reserva Forestal San Ram&#243;n, Cordillera de Tilar&#225;n, Los &#193;ngeles, banks of the road between Los &#193;ngeles and R&#237;o Cataratas, 10˚12'15''N, 84˚32'15''W, 900 - 1250 m, 2 May 1996, A. Rojas &amp; J. Morales 2589 (CR). Cartago: Pacayas, near of bridge over the R&#237;o Turrialba, trail to Turrialba (between Santa Teresa and Santa Cruz), 9˚57'20''N, 83˚46'20''W, 1670 m, 27 Oct 1995, A. Cascante et al. 847 (CR); 10 km S of Cartago by air, along confluence of r&#237;o Empalme and R&#237;o Estrella, 1 km S of Palo Verde by road, Palo Verde is 1 km S of Panamerican hwy on road to La Estrella, 9˚46'N, 83˚57'W, 1450 m, 21 Apr 1983, R. Liesner &amp; E. Judziewicz 14587 (MO); Para&#237;so, Cordillera de Talamanca, Estaci&#243;n de Biolog&#237;a Tropical R&#237;o Macho and surroundings, Orosi, 09˚46'N, 83˚52'W, 1550 - 1580 m, 30 Sep 1993, A. Rojas 480 (CR, INB, MO); Guadalupe (Arenilla), Zona Protectora La Carpintera, slope of hill of La Carpintera, with view to Coris, 9˚52'39.2''N, 83˚58'17.6''W, 1710 m, 19 Mar 2007, A. Ruiz et al. 1179 (CR); 0.8 km down (W) road to San Cristobal Norte, 1850 m, 21 Set 1979, K. Walter 79498 (CR). Guanacaste: La Cruz, Parque Nacional Guanacaste, 9 km S of Santa Cecilia, Estaci&#243;n Pitilla, 10˚59'26''N, 85˚25'40''W, 700 m, 28 Sep 1990, C. Ch&#225;vez et al. 188 (CR, MO); Monteverde, 3.4 km N of Santa Elena on road to Las Nubes, 10˚22ʹN, 84˚49ʹW, 1500 m, 14 Aug 1988, W. Haber &amp; W. Zuchowski 8547 (CR, MO); Monteverde, 3.5 km N of Santa Elena on road to San Gerardo, 0.5 km N of junction road and R&#237;o Negro, 10˚21'N, 84˚48'W, 1540 m, 20 Aug 1988, W. Haber &amp; W. Zuchowski 8579 (CR); Cordillera de Tilar&#225;n, 1 km N of Las Nubes de R&#237;o Chiquito, Monteverde zone, Atlantic slope, 10˚22'N, 84˚51'W, 1300 m, 4 Sep 1988, W. Haber &amp; W. Zuchowski 8689 (CR, MO); Tilar&#225;n, Cordillera de Tilar&#225;n, R&#237;o San Lucas, 10˚22'50''N, 84˚54'20''W, 800 - 900 m, 27 Jun 1995, G. Rodr&#237;guez &amp; A. Rojas 357 (CR); La Uni&#243;n, San Rafael, Zona Protectora Cerros La Carpintera, Iztar&#250; camp, between the road and the top of the mountain, 9˚53'11''N, 83˚58'16''W, 1750 - 1850 m, 17 Jul 2007, A. Rojas et al. 7599 (CR). Heredia: Vara Blanca, 1700 - 1900 m, 27 Oct 1979, C. Todzia 929 (CR). Lim&#243;n: Talamanca, trail ca. 5 km up Alto Lari, 9˚24'15''N, 83˚05'05''W, 1300 m, 27 Feb 1992, J. Bittner 1338 (CR); Talamanca, Bratsi, Amubri, Alto Lari, Kivut, 9˚24'15''N, 83˚05'15''W, 1300 m, 6 Mar 1992, G. Herrera 5174 (CR). Puntarenas: Coto Brus, Parque Internacional La Amistad, Cordillera de Talamanca, Estaci&#243;n Pittier, R&#237;o Gemelo trail, 9˚01'30''N, 82˚57'40''W, 1680 m, 29 Jan 1995, F. Alvarado et al. 12 (INB, CR); upper r&#237;o Bur&#250;, 2010 m, 19 Aug 1983, L. G&#243;mez et al. 21431 (CR); Ojo de Agua, Monteverde, M&#233;ndez estate, R&#237;o Aranjuez, 10˚17'N, 84˚46'W, 1550 m, 14 Nov 1987, W. Haber &amp; E. Bello 7731a (CR); La Pitahaya, Arancibia, Rinc&#243;n, headers of R&#237;o Aranjuez, 10˚15'10''N, 84˚41'20''W, 1200 m, 27 May 1996, G. Herrera 9024 (CR); Monteverde Reserve, 1 km SW station, 10˚18'N, 84˚48'W, 1500 - 1550 m, 20 Feb 1992, S. Ingram &amp; K. Ingram 1361 (CR); Monteverde, 1520 m, 13 Nov 1979, S. Koptur 223 (CR); Monteverde, 1 Dec 1979, S. Koptur 246 (CR); Monteverde, Cordillera de Tilar&#225;n, 1350 m, 30 Jul 1981, D. Neill 5067 (CR); Monteverde, Cordillera de Tilar&#225;n, 4500 ft [=1372 m], Mar 1959, C. Palmer 110 (CR); Coto Brus, Zona Protectora Tablas, Cordillera de Talamanca, Sabalito, Las Alturas de Cot&#243;n, Estaci&#243;n Biol&#243;gica Las Alturas, surroundings of the station, 8˚57'15''N, 82˚50'10''W, 1580 m, 23 Dec 1993, A. Rojas 765 (INB, MO); Monteverde Cloud Forest Reserve, road to station from comunity, 10˚06ʹN, 83˚26'W, 1500 m, 23 Aug 1987, J. Smith 563 (CR). San Jos&#233;: P&#233;rez Zeled&#243;n, Cordillera de Talamanca, Las Nubes de Santa Elena, 9˚23'30''N, 83˚35'50''W, 1210 m, 8 sep 1995, E. Alfaro 395 (CR, INB); Desamparados, Patarr&#225;, Cerro El Espino (Mata de Ca&#241;a-Azahar), 1600 - 1800 m, 13 Nov 1983, I. Chac&#243;n &amp; G. Herrera 1564 (CR); ib&#237;dem, I. Chac&#243;n &amp; G. Herrera 1569 (CR); Coronado, Las Nubes, 29 Dec 1956, F. Chisaki 1004 (CR); surroudings S of San Jos&#233;, trail more than 1 km NE of Tarbaca to Joya, which deviates from the road no. 4 Aserr&#237;-Tarbaca,</p><p>1750 m, 29 Apr 1990, P. D&#246;bbeler 112 (MEXU); Tarraz&#250;, San Carlos, trail to San Josecito, 9˚37'20''N, 84˚07'30''W, 1400 m, 23 set 1997, A. Estrada 1177 (CR); Escaz&#250;, cerros de Escaz&#250;, to Alto Tapezco by Bebedero, 9˚55'00''N, 84˚10'00''W, 1100 - 1700 m, 3 Oct 1993, B. Hammel et al. 19070 (CR); Dota, Peor es Nada, NW from the foothills of Cerro Lira, 9˚31'30''N, 83˚51'40''W, 1800 m, 8 Dec 1994, G. Herrera &amp; S. Mart&#233;n 7410 (CR); Dota, Copey, Providencia, Zapotal, path on the right after the river, 9˚31'50''N, 83˚49'49''W, 1800 m, 20 Aug 2003, S. Lobo &amp; A. Ruiz 507 (CR); Aserr&#237;, Cerros de Escaz&#250;, Tarbaca, 9˚49'10''N, 84˚07'30''W, 1850 m, 12 Oct 1993, J. Morales 1871 (CR); Parque Nacional Braulio Carrillo, ca. 1 km along road from entrance, 1500 m, 19 Jul 1983, R. Moran 3294 (CR, F); Dota, Copey, Zapotal de Providencia, of the Monta&#241;a Fr&#237;a farm 0 - 1 km to Providencia, 9˚32'05''N, 83˚50'19''W, 1720 m, 26 Aug 2004, A. Rojas &amp; H. G&#243;mez 5975 (CR, MO); Tarraz&#250;, San Carlos, Cerro Cura, trail to San Marcos, 9˚36'17''N, 84˚06'39''W, 1730 m, 21 Sep 2004, A. Rojas &amp; S. Lobo 6096 (CR); Aserr&#237;, Cerros de Escaz&#250;-La Carpintera, Tarbaca, El Cedral, trail to Pico Alto, 9˚49'35''N, 84˚06'50''W, 1770 - 2000 m, 16 Jun 1996, A. Rojas &amp; M. Obando 2628 (CR); Acosta, Palmichal, Zona Protectora Cerros de Escaz&#250;, side of R&#237;o Tabarcia, 9˚50'32''N, 84˚10'21''W, 1450 m, 23 Jun 2004, A. Rojas &amp; A. Quesada 5789 (CR); along R&#237;o Zurqu&#237;, (downstream from Gu&#225;piles hwy), Cordillera Central, 10˚03'N, 84˚0'W, ca. 1400 - 1500 m, 18 Jan 1986, A. Smith et al. 1731 (CR, UC); Escaz&#250;, Piedra Blanca, 1900 m, 20 Jan 1935, F. Sol&#237;s 149 (CR); San Vicente, La Justa, 1170 m, Sep 1925, M. Valerio 124 (CR).</p><p>Campyloneurum leoniae separates from C. irregulare because has not pruinose rhizome (vs. pruinose); fronds approximate to 3 (-5) mm distant (vs. (approximate-) 2 - 5 mm distant); rhizome scales with the apical cells 2 - 5 timers longer tan wide (vs. 1.5 - 3.5 times); phyllopodia relatively shorter (0.5 - 1 (-2) mm vs. 1 - 3 mm); adaxial blade surface opaque (vs. lustrous (rarely opaque)) and distributed to lower elevation ((700-) 1000 - 1850 (-2000) m vs. (1700-) 190 - 2500 m) (See <xref ref-type="fig" rid="fig2">Figure 2</xref> and <xref ref-type="table" rid="table2">Table 2</xref>).</p><fig id="fig2"  position="float"><label><xref ref-type="fig" rid="fig2">Figure 2</xref></label><caption><title> (a) (b). Type material of Campyloneurum leoniae A. Rojas (W. Haber &amp; W. Zuchowski 8691, CR). (a) Habit; (b) Rhizome scales; (c) Rhizome scales of Campyloneurum irregulare Lellinger (A. Rojas et al. 6229, CR)</title></caption><graphic mimetype="image"   position="float"  xlink:type="simple"  xlink:href="http://html.scirp.org/file/19-2603096x3.png"/></fig><table-wrap id="table2" ><label><xref ref-type="table" rid="table2">Table 2</xref></label><caption><title> Comparison of morphological characteristics among four species of Campyloneurum: C. amphostenon, C. irregulare, C. leoniae and C. pittieri</title></caption><table><tbody><thead><tr><th align="center" valign="middle" >Character</th><th align="center" valign="middle" >C. amphostenon</th><th align="center" valign="middle" >C. irregulare</th><th align="center" valign="middle" >C. leoniae</th><th align="center" valign="middle" >C. pittieri</th></tr></thead><tr><td align="center" valign="middle" >Rhizome glaucous</td><td align="center" valign="middle" >yes</td><td align="center" valign="middle" >yes</td><td align="center" valign="middle" >no</td><td align="center" valign="middle" >No or scarcely</td></tr><tr><td align="center" valign="middle" >Distance between fronds</td><td align="center" valign="middle" >2 - 15 mm</td><td align="center" valign="middle" >(approximate-) 2 - 5 mm distant</td><td align="center" valign="middle" >Approximate to 3 (-5) mm distant</td><td align="center" valign="middle" >5 - 30 (-50) mm</td></tr><tr><td align="center" valign="middle" >Long of rhizome scales</td><td align="center" valign="middle" >4 - 8 mm</td><td align="center" valign="middle" >3 - 5 mm</td><td align="center" valign="middle" >1.5 - 4.0 mm</td><td align="center" valign="middle" >2 - 5 mm</td></tr><tr><td align="center" valign="middle" >Color of rhizome scales</td><td align="center" valign="middle" >yellowish-brown to grayish-brown</td><td align="center" valign="middle" >brown to dark brown</td><td align="center" valign="middle" >brown to dark brown</td><td align="center" valign="middle" >brown to dark brown</td></tr><tr><td align="center" valign="middle" >Form of basal cell of rhizome scales</td><td align="center" valign="middle" >triangular to hexagonal, isodiametric</td><td align="center" valign="middle" >poligonal, irregular</td><td align="center" valign="middle" >poligonal, irregular</td><td align="center" valign="middle" >Rounded to poligonal, isodiametric</td></tr><tr><td align="center" valign="middle" >Long of apical cells of rhizome scales</td><td align="center" valign="middle" >1.5 - 3.0 times longer than wide</td><td align="center" valign="middle" >1.5 - 3.5 times longer than wide</td><td align="center" valign="middle" >2 - 5 times longer than wide</td><td align="center" valign="middle" >1 - 2 times longer than wide</td></tr><tr><td align="center" valign="middle" >Lumen cells color of rhizome scales</td><td align="center" valign="middle" >translucid</td><td align="center" valign="middle" >brown</td><td align="center" valign="middle" >brown</td><td align="center" valign="middle" >translucid to grayish-brown</td></tr><tr><td align="center" valign="middle" >Fronds size</td><td align="center" valign="middle" >30 - 55 (-65) cm</td><td align="center" valign="middle" >20 - 45 cm</td><td align="center" valign="middle" >(10-) 20 - 35 (-50) cm</td><td align="center" valign="middle" >(35-) 45 - 60 (-80) cm</td></tr><tr><td align="center" valign="middle" >Long of phyllopodia</td><td align="center" valign="middle" >1.5 - 3 mm</td><td align="center" valign="middle" >1 - 3 mm</td><td align="center" valign="middle" >0.5 - 1 (-2) mm</td><td align="center" valign="middle" >2 - 4 mm</td></tr><tr><td align="center" valign="middle" >Long of stipe</td><td align="center" valign="middle" >4 - 10 (-20) cm</td><td align="center" valign="middle" >0.5 - 3.0 (-5.0) cm</td><td align="center" valign="middle" >(0.5-) 1 - 5.0 cm</td><td align="center" valign="middle" >8 - 25 cm</td></tr><tr><td align="center" valign="middle" >Diameter of stipe</td><td align="center" valign="middle" >2 - 3 mm</td><td align="center" valign="middle" >(1-) 1.5 - 2 mm</td><td align="center" valign="middle" >1 - 1.5 mm</td><td align="center" valign="middle" >2 - 3 mm</td></tr><tr><td align="center" valign="middle" >Broad of blade</td><td align="center" valign="middle" >1.0 - 3.5 cm</td><td align="center" valign="middle" >(0.4-) 0.6 - 1.2 cm</td><td align="center" valign="middle" >(0.5-) 1 - 2 (-.3) cm</td><td align="center" valign="middle" >(1.3-) 2.0 - 5.0 cm</td></tr><tr><td align="center" valign="middle" >Brightness of abaxial blade surface</td><td align="center" valign="middle" >middle to markerly lustrous</td><td align="center" valign="middle" >lustrous (rarely opaque)</td><td align="center" valign="middle" >opaque</td><td align="center" valign="middle" >middle lustrous</td></tr><tr><td align="center" valign="middle" >Highlighted veins</td><td align="center" valign="middle" >few or not</td><td align="center" valign="middle" >not</td><td align="center" valign="middle" >not</td><td align="center" valign="middle" >yes</td></tr><tr><td align="center" valign="middle" >Altitudinal distribution</td><td align="center" valign="middle" >(1800-) 2100 - 2900 (-3200) m</td><td align="center" valign="middle" >(1700-) 1900 - 2500 m</td><td align="center" valign="middle" >(700-) 1000 - 1850 (-2000) m</td><td align="center" valign="middle" >(1900-) 2700 - 3400 (-3550) m</td></tr></tbody></table></table-wrap></sec><sec id="s3_1_3"><title>3.1.3. Campyloneurum madrense A. Rojas, Sp. Nov. (<xref ref-type="fig" rid="fig3">Figure 3</xref>).</title><p>TYPE: MEXICO. Chiapas: Mpio. Ocosingo, 6.35 km NW of San Javier cruise, 16˚50'34''N, 91˚23'W, 391 m, 15 Nov 2003, D. &#193;lvarez et al. 6960 (holotype: MEXU).</p><p>Diagnoses: Campyloneurum madrense differs from C. angustifolium by not pruinose rhizome or scarcely so, linear rhizome scales with sparsely denticulate margin, they dark gray to blackish, inside with narrow oblong cells and distributed at lower altitudinal distribution.</p><p>Description: Epiphytic; rhizome 2 - 5 mm in diameter, compact to short-creeping, fronds approximate to distant to 2 mm, not pruinose, brown to blackish when dry; rhizome scales 3 - 6 &#215; 0.4 - 0.8 mm, linear to linear-lanceolate, dark gray to blackish, clathrate, dense, apically long acute to attenuate, marginally sparsely denticulate, cells oblong to narrowly oblong, 2 - 5 times longer than wide, lumen gray; fronds (18-) 30 - 65 cm long; stipe 2 - 5 (-8) cm long, 1/12 - 1/8 of the frond length, 1 - 1.5 mm wide (to 2 mm with wing), slight to strongly flattened, strawish to pale brown, winged throughout, glabrous or sparsely scaly, the scales 0.5 - 1.5 x ca. 0.5 mm, ovate, brown; phyllopodia 0.5 - 1 (-2) mm long; blade (16-) 27 - 48 &#215; (0.5-) 0.8 - 1.5 cm, linear, attenuate on both sides, chartaceous to subcoriaceous, opaque to lustrous, with commonly revolute margin; rachis strawish to brown, glabrous or sparsely scaly at base; lateral veins few evident; sori rounded, in 1 - 2 (-3) irregular rows between costa and margin.</p><p>Etimology: The name of it species make reference to geographical distribution in Sierra Madre Oriental and Sierra Madre del Sur.</p><p>Distribution: Actually known only from Mexico at 25 - 940 m elevation.</p><p>Additional revised specimens (paratypes): MEXICO. Chiapas: Municipio Ocosingo, 2 km S of Nuevo Guerrero, 16˚59'02''N, 91˚17'07''W, 248 m, 31 Jan 2003, G. Aguilar 5371 (MEXU); Municipio Ocosingo, 0.83 km SE of Nuevo Guerrero village, 16˚58'51''N, 91˚16'44''W, 222 m, 4 Nov 2003, G. Aguilar 8347 (MEXU); Municipio Ocosingo, 4.2 km E of Nuevo Francisco Le&#243;n, 16˚19'43''N, 91˚16'57''W, 180 m, 13 Oct 2002, G. Aguilar &amp; D. &#193;lvarez 3443 (MEXU); Municipio Ocosingo, 6.37 km SE of Nuevo Guerrero, trail of Lancanj&#225;-Tzeltal, 16˚55'57''N, 91˚15'43''W, 456 m, 15 Oct 2002, G. Aguilar &amp; D. &#193;lvarez 3575 (MEXU) Municipio Ocosingo, 1.3 km S of El Para&#237;so, 16˚57'05''N, 91˚15'43''W, 483 m, 16 Oct 2002, G. Aguilar &amp; D. &#193;lvarez 3642 (MEXU); Municipio Ocosingo, 0.09 km SO of Lacanj&#225; Chansayab community, 16˚45'39''N, 91˚07ʹ47ʺW, 325 m, 15 Nov 2003, G. Aguilar &amp; C. Chancayun 8425 (MEXU); Municipio Ocosingo, waterhole of San Javier, 16˚50'25''N, 91˚08'32''W, 390 m, 2 Dec 2002, G. Aguilar &amp; A. Cort&#233;s 4504 (MEXU); Municipio Ocosingo, Chansayab community, 16˚45'16''N, 91˚07'55''W, 362 m, 19 Oct 2002, G. Aguilar et al. 3830 (MEXU), ib&#237;dem, G. Aguilar et al. 3832 (MEXU); Municipio Ocosingo, 4.8 km SE of Nuevo Guerrero, 16˚57'26''N, 81˚15'02''W, 338 m, 23 May 2002, D. &#193;lvarez &amp; G. Aguilar 1420 (MEXU); Municipio Ocosingo, 4.5 km SE of Bonampak archaeological zone, 16˚40'33''N, 91˚02'05''W, 256 m, 10 Feb 2003, D. &#193;lvarez &amp; A. Chambor 3720 (MEXU); Municipio Ocosingo, 7.5 km SW of Bonampak, 16˚39'51''N, 91˚00'59''W, 25 m, 28 Nov 2002, D. &#193;lvarez et al. 2743 (MEXU) Municipio Ocosingo, 6,35 km NW of the cruise of San Javier, 16˚34ʹN, 91˚08'43''W, 391 m, 15 Nov 2003, D. &#193;lvarez et al. 6960 (MEXU); Municipio Ocozocoautla de Espinoza, 26 - 28 km N of Ocozocoautla, along road to Mal Paso, 700 m, 15 Nov 1971, D. Breedlove &amp; A. Smith 22445 (MEXU); 13 mi N of Ocozocoautla, on gravel road to Apitpac, near km 21 marker, 900 m, 9 Jul 1977, T. Croat 40605 (MEXU, MO); along dirt road between San Fernando and Moravillas (near Lago Malpaso), 4 - 66 mi NW of San Fernando, 16˚53'N, 93˚16'W, 840 - 940 m, 15 Feb 1987, T. Croat &amp; D. Hannon 65024 (MEXU, MO); along hwy 197, between Chiapa de Corzo and Pichucalco, 0.6 mi N of Tapulula, 35.6 mi SSE of Pichucalco, vicinity of km 124 marker, 17˚16'N, 93˚01'W, 750 m, 18 Feb 1987, T. Croat &amp; D. Hannon 65272 (MEXU, MO); Municipio Ocosingo, 4 km SE of Nuevo Guerrero, trail to Boca Lacantum, 560 m, 24 Jan 1986, E. Mart&#237;nez 16764 (MEXU); Municipio Ocozocoautla, Reserva Ecol&#243;gica “El Ocote”, 750 m, 17 Feb 1986, M. Palacios 2884 (MEXU, UC, XAL); Municipio Ocosingo, R&#237;o Cedro, W of Lacanja Chansayab, 300 m, 15 Jan 1984, F. V&#225;zquez &amp; S. Avenda&#241;o 1514 (MEXU, XAL). Oaxaca: Municipio San Pedro Ixcatlan, In the village of Cerro Quemado, road entrance Ojitl&#225;n-Xalapa de Diaz, 230 m, 8 Feb 1984, J. Calzada 10263 (UC, XAL); Municipio Soyaltepec, Distrito Tuxtepec, spillway of the Temascal dam, 18˚12'N, 96˚20'W, 70 m, 21 oct 1987, L. Cortes &amp; R. Torres 1092 (MEXU); along road from Valle Nacional to Arriba Tortuga and Armadillo, via Rancho Grande and Loma San Rafael, 5 - 6 km NW of Valle Nacional Center (Hotel del Valle), 17˚47'25''N, 96˚17'55''W, 474 - 476 m, 2 Mar 2008, T. Croat 100128 (MO, XAL); Municipio Santa Mar&#237;a Chimalapa, near of Arroyo Sangre, ca. 2 km E de Santa Mar&#237;a, 16˚54'30''N, 94˚40'30''W, 280 m, 27 Aug 1984, H. Hern&#225;ndez 369 (CHAPA, MEXU); Chiltepec, Tuxtepec, 7 Aug 1965, G. Mart&#237;nez 174 (MEXU); Sierra de Ju&#225;rez, Municipio Valle Nacional, Yetla, km 154 of trail Tuxtepec-Oaxaca, 800 m, 15 Jun 1966, G. Mart&#237;nez 879 (MEXU, XAL); Distrito Tuxtepec, Municipio Valle Nacional, along hwy M&#233;xico 175, 21 km (by road) N of Vista Hermosa and 4 km SW of Valle Nacional, 17˚44'N, 96˚21'W, 450 m, 15 Dec 1985, M. Nee 32158 (NY, UC, XAL); Municipio Putla, Distrito Putla, ca. 4 km NE of San Pedro, 800 m, 11 Jul 1988, E. Solano 429 (CHAPA); Municipio Santa Mar&#237;a Jacatepec, Distrito Tuxtepec, trail to Cosolapa San Antonio, Ejido San Felipe Tilpa, 13.3 km SW of La Reforma, 17˚51'N, 96˚03'W, 300 m, 20 Feb 1988, R. Torres &amp; L. Cortes 11439 (MEXU); Municipio Mat&#237;as Romero, hills between R&#237;o Verde and Arroyo Hamaca, near R&#237;o Verde, 10 km SE from the La Foresta sawmill (abandoned), ca. 24 km S of Esmeralda, 17˚01'N, 94˚44'W, 300 m, 27 Sep 1982, T. Went et al. 4007 (CHAPA, UC); Santa Mar&#237;a Chimalapa, Arroyo Sangre, ca. 1 - 2 km W of Santa Mar&#237;a, 16˚54'30''N, 94˚40'W, 250 m, 27 Jul 1985, T. Went et al. 5132 (MEXU). Puebla: Municipio Hueytamalco, 1 km NE of Campo Experimental “Las Margaritas” facilities, Instituto Nacional de Investigaciones Forestales, Agr&#237;colas y Pecuarias (INIFAP), 20˚00'25''N, 97˚18'45''W, 490 m, 16 Jun 2007, G. Ibarra et al. 5333 (MEXU). Tabasco: Municipio Tacotalpa, Cerro del Madrigal, km 7 of Tacotalpa station to Tapijulapa, 13 Apr 1980, C. Cowan 2908 (MEXU); 13 mi N of Ocozocuautla on gravel road to Apitac, near km 21 marker, 900 m, 9 Jul 1977, T. Croat 40619 (MEXU, MO); Municipio Teapa, surroundings of Centro Regional Puyacatengo, 16 Jan 1982, S. Zamudio 180 (XAL). Veracruz: Municipio Atoyac, Ejido Esperanza, 18˚52'00''N, 96˚44'00''W, 750 m, 21 Jun 1985, R. Acevedo &amp; G. Castillo 281 (XAL); Municipio Atoyac, Miraflores, 18˚57'00''N, 96˚49'00''W, 750 m, 22 Jun 1985, R. Acevedo &amp; G. Castillo 286 (MEXU); Municipio San Andr&#233;s Tuxtla, Estaci&#243;n de Biolog&#237;a Tropical Los Tuxtlas, 150 m, 5 Oct 1974, J. Calzada 1547 (XAL); Municipio Catemaco, hills S of Cerro Jegal, N of Los Naranjos Ranch, road entrance Catemaco-Dos Amates, 21 May 1985, J. Calzada 11575 (XAL); Municipio Soteapan, Ejido Piedra Labrada to N, 17 May 1985, J. Calzada et al. 11518 (XAL); Municipio Catemaco, El Naranjillo, El Naranjillo Ranch, road Catemaco-Sontecomapan, 18˚25'N, 95˚06'W, 7 Aug 1993, G. Carmona 60 (XAL); Municipio San Andr&#233;s Tuxtla, El Bastonal, 18 Sep 1985, R. Cedillo 3402 (MEXU, XAL); Municipio Jes&#250;s Carranza, km 6 from trail Cedillo-R&#237;o Alegre, 17˚10'N, 94˚40'W, 150 m, 18 Jan 1975, J. Dorantes 3916 (XAL); Municipio Catemaco, Coyame, near Catemaco, 18˚27'N, 95˚05'W, 180 m, 22 Jan 1972, R. Hern&#225;ndez 1386 (MEXU, XAL); Municipio Catemaco, Ejido Balzapote, ca. 3 km N of the Los Tuxtlas Biological Station, 15 Oct 1992, P. Hietz &amp; U. Seifert 562 (MEXU); Municipio San Andr&#233;s Tuxtla, Laguna Escondida, 5 km NE of Estaci&#243;n de Biolog&#237;a Tropical Los Tuxtlas, 18˚34 - 36'N, 95˚04 - 09'W, 250 m, 12 Sep 1982, G. Ibarra 307 (MEXU); Municipio Montepio, Estaci&#243;n de Biolog&#237;a Tropical Los Tuxtlas, 18˚34'N, 94˚05'W, ca. 140 m, 29 Jul 1976, H. Kennedy &amp; C. Horvitz 3676 (MEXU); 33 km NE of Catemaco, Biological Station, 200 m, 30 Apr 1973, K. Kramer 1974 (MEXU); Municipio San Andr&#233;s Tuxtla, Estaci&#243;n de Biolog&#237;a Tropical Los Tuxtlas, Cerro Vigia 5, lot 67, 18˚34'N, 95˚04'W, 450 m, 12 Apr 2005, T. Kr&#246;mer et al. 1924 (MEXU); Municipio San Andr&#233;s Tuxtla, Colegio Agr&#237;cola Militar, Montepio “Rancho Pouchoulen”, 18˚34'N, 95˚08'W, 740 m, 28 Apr 2005, T. Kr&#246;mer et al. 2029 (MEXU, XAL); Municipio Catemaco, La Palma, Don Fidel plot, 18˚33'N, 95˚04'W, 35 m, 8 Aug 2005, T. Kr&#246;mer et al. 2465 (MEXU); Municipio Catemaco, 5 km E of Tebanca course to Bastonal, N slope of Sierra de Santa Marta, 830 m, 10 Dec 1980, R. Lira 170 (XAL); Municipio Las Choapas, “El Milagro” Ranch, 5 km in a straight line to SW (202˚) of Nueva Tabasque&#241;a colony, 17˚31'48''N, 94˚01'44''W, 115 m, 27 Aug 2002, E. L&#243;pez 241b (XAL); Municipio San Andr&#233;s Tuxtla, Estaci&#243;n de Biolog&#237;a Tropical Los Tuxtlas, between Montepio and Sontecomapan, 150 m, 12 Jun 1981, D. Lorence 3477 (XAL); Municipio Catemaco, ca. 8 km after Coyame course to Adolfo L&#243;pez Mateos, 18˚26'22''N, 95˚00'11''W, 650 m, 3 May 2002, A. Mendoza et al. 617b (MEXU); Municipio Pajapan, 5 km NW of Pajapan, SE slopes of Cerro San Mart&#237;n Pajapan, 18˚17'30''N, 94˚43'W, 700 m, 3 Nov 1981, M. Nee &amp; J. Calzada 22768 (MEXU, XAL); gravel Cliff 1 mi W of Fort&#237;n, 4 Aug 1947, J. Paxson et al. 17M680 (MEXU, UC); Municipio San Andr&#233;s Tuxtla, Colegio Agr&#237;cola Militar, Montepio, Huber Ranch, 18˚36'N, 95˚05'W, 70 m, 7 Sep 2005, A. P&#233;rez et al. 11 (MEXU); Municipio San Andr&#233;s Tuxtla, Estaci&#243;n de Biolog&#237;a Tropical Los Tuxtlas, 28 Oct 1981, T. Ramamoorthy 2613 (MEXU); Municipio San Andr&#233;s Tuxtla, Estaci&#243;n de Biolog&#237;a Tropical Los Tuxtlas, 19 Jan 1982, T. Ramamoorthy &amp; G. Ibarra 3268 (MEXU); Municipio Catemaco, Pen&#237;nsula de Moreno, tral Coyame-Pen&#237;nsula de Moreno, 18˚27'N, 95˚00'W, 200 m, 18 Aug 1980, F. Ram&#237;rez 1382 (XAL); Municipio Catemaco, trail Tebanca-El Bastonal, 18˚24'N, 94˚59'W, 400 m, 27 mar 1980, F. Ram&#237;rez &amp; F. V&#225;zquez 704 (XAL); Municipio Catemaco, Santa Rosa, 18 km S of Catemaco, 450 m, 10 Apr 1973, R. Riba et al. 705 (MEXU); San Pedro Soteapan, Arroyo Soteapan, 1 km W ofSan Fernando, S slope of Sierra de Santa Marta, 600 m, 13 Aug 1980, R. Riba et al. 1135 (MEXU); Municipio Catemaco, Pen&#237;nsula de Moreno, ca. 4 km E of Coyame, 300 m, 17 ago 1980, R. Riba et al. 1223 (MEXU); Municipio Coetzala, hillside E of Coetzala, 18˚46'50''N, 96˚54'52''W, 650 m, 4 Nov 2001, A. Rinc&#243;n &amp; C. Dur&#225;n 2819 (XAL); Municipio Catemaco, Cerro Jegal, 8 km N of Catemaco, 18˚29'05''N,</p><p>95˚03'53''W, 940 m, 4 nov 2000, A. Rinc&#243;n et al. 2137 (MEXU, XAL); some km before Montepio, 18˚38'N, 95˚05'W, 2 May 1988, J. van Rooden 810 (MEXU); Municipio Catemaco, Dos Amates de Sontecomapan, 4 Aug 1965, 500 m, J. Rzedowski 20356 (MEXU); Municipio Amatl&#225;n, Cerro Cruztitl&#225;n, near the cave chapel of La Virgen, in front of Pe&#241;uelas, E of Amatl&#225;n, 18.84704˚N, 96.88715˚W, 743 m, 30 Jan 2015, G. Salazar et al. 9376 (MEXU); Municipio Catemaco, at highest point on road from Catemaco to Sontecomapan, 5 km N of juntion with road around Lago Catemaco, 8 km (by air) NE of Catemaco, 18˚29'N, 95˚04'W, 510 m, 14 Jan 1981, G. Schatz &amp; M. Nee 227 (XAL); Municipio Catemaco, Dos Amates, Cerro Chochobi, 720 m, 20 Dec 2006, B. Senterre &amp; M. Parvais 4379 (XAL); Municipio San Andr&#233;s Tuxtla, Estaci&#243;n de Biolog&#237;a Tropical Los Tuxtlas, 18˚34'N - 18˚36'N, 95˚04W - 95˚09'W, 700 m, 10 Sep 1986, S. Sinaca 921 (MEXU); Municipio San Andr&#233;s Tuxtla, Ejido Balzapote, 5 km N of Estaci&#243;n de Biolog&#237;a Tropical Los Tuxtlas, 18˚34'N - 18˚36'N, 95˚04'W - 95˚09'W, 180 m, 16 Feb 1989, S. Sinaca &amp; P. Ventura 1490 (MEXU); 3 km N of Catemaco, along dirt road, 18˚30'N, 95˚10'W, 400 m, 15 Jun 1982, S. Solheim &amp; V. Powers 848 (XAL); Municipio Catemaco, Estaci&#243;n de Biolog&#237;a Universidad de Veracruz, 500 - 650 m, 14 Sep 1998, A. Torrez &amp; A. Campos 18 (MEXU); Municipio Hidalgotitl&#225;n, E of Campamento Hermanos Cedillo to La Laguna, 1 km of camp, 11 Apr 1974, P. Valdivia 212 (XAL); Municipio Hidalgotitl&#225;n, 2, 5 km of Campamento Hermanos Cedillo in front of Paso Moral, N side, 17˚15'30''N, 94˚36'W, 150 m, 18 Apr 1974, P. Valdivia 324 (XAL); Municipio Hidalgotitl&#225;n, to &#193;lvaro Obreg&#243;n, 6 km of Campamento Hermanos Cedillo, 17˚18.7'N, 94˚39'W, 149 m, 23 Apr 1974, P. Valdivia 416 (XAL); Municipio Hidalgotitl&#225;n, hacia &#193;lvaro Obreg&#243;n, 6 km of Campamento Hermanos Cedillo, 17˚18,7'N, 94˚39'W, 150 m, 23 Apr 1974, P. Valdivia 426 (XAL); Municipio Hidalgotitl&#225;n, to &#193;lvaro Obreg&#243;n, 300 m of Campamento Hermanos Cedillo, by Plan de Arroyo, W direction, 17˚16'N, 94˚37'W, 149 m, 24 Apr 1974, P. Valdivia 452 (XAL); Municipio Hidalgotitl&#225;n, 6.5 km of Campamento Hermanos Cedillo to La Laguna, 17˚16.6'N, 94˚33.4'W, 159 m, 23 May 1974, P. Valdivia 615 (XAL); Municipio Hidalgotitl&#225;n, in km 6.06 and to 200 m of N side course to La Laguna, 17˚16.3'N, 94˚35.2'W, 160 m, 28 May 1974, P. Valdivia 760 (XAL); Municipio Hidalgotitl&#225;n, 1.7 km of Campamento Hermanos Cedillo to La Escuadra, 17˚17'N, 94˚36.5'W, 156 m, 22 Jun 1974, P. Valdivia 915 (XAL); Municipio Hidalgotitl&#225;n, 1.5 km of Campamento Hermanos Cedillo to La Escuadra, 17˚16.5'N, 94˚35.3'W, 156 m, 24 Jun 1974, P. Valdivia 943 (XAL); Municipio Hidalgotitl&#225;n, 800 m of Campamento Hermanos Cedillo to La Escuadra, 17˚15'N, 94˚37'W, 156 m, 22 Jul 1974, P. Valdivia 995 (XAL), ib&#237;dem, 24 Jul 1974, P. Valdivia 1075 (XAL); Municipio Hidalgotitl&#225;n, 1 km of Campamento Hermanos Cedillo to La Escuadra, 17˚15'N, 94˚37'W, 156 m, 29 Jul 1974, P. Valdivia 1164 (XAL); Municipio Hidalgotitl&#225;n, 2 km of Campamento Hermanos Cedillo to La Escuadra, 17˚17.3'N, 94˚36.5'W, 156 m, 31 Jul 1974, P. Valdivia 1202 (XAL), ib&#237;dem, 01 Aug 1974, P. Valdivia 1229 (XAL),</p><p>ib&#237;dem, P. Valdivia 1242 (XAL); Municipio Hidalgotitl&#225;n, 1.5 km of Campamento Hermanos Cedillo to La Escuadra, 17˚15.5'N, 94˚37'W, 150 m, 2 Aug 1974, P. Valdivia 1259 (XAL); Municipio Hidalgotitl&#225;n, 6 km of Campamento Hermanos Cedillo to La Escuadra, 17˚18,7'N, 94˚39'W, 150 m, 06 Aug 1974, P. Valdivia 1292 (XAL); Municipio Hidalgotitl&#225;n, La Escuadra, 17˚19'N, 94˚38.5'W, 160 m, 07 Aug 1974, P. Valdivia 1325 (XAL); Municipio Hidalgotitl&#225;n, 500 m of Campamento Hermanos Cedillo to La Escuadra, 17˚14'N, 94˚37'W, 156 m, 09 Oct 1974, P. Valdivia 1380 (XAL); Municipio Hidalgotitl&#225;n, 6 km of Campamento Hermanos Cedillo to La Laguna, 17˚16'N, 94˚32.5'W, 156 m, 12 Aug 1974, P. Valdivia 1440 (XAL); Municipio Hidalgotitl&#225;n, 500 m of Campamento Hermanos Cedillo to La Escuadra, 17˚16.5'N, 94˚37.5'W, 156 m, 14 ago 1974, P. Valdivia 1469 (XAL); Municipio Hidalgotitl&#225;n, a 1.5 km of Campamento Hermanos Cedillo to La Escuadra, 17˚15.5'N, 94˚37'W, 156 m, 13 Dec 1974, P. Valdivia 1600 (XAL); Municipio Hidalgotitl&#225;n, 4.5 km of Campamento Hermanos Cedillo to La Escuadra, 17˚16.5'N, 94˚38'W, 149 m, 19 Dec 1974, P. Valdivia 1706 (XAL); Municipio Hidalgotitl&#225;n, 3 km of Campamento Hermanos Cedillo to La Escuadra, 17˚16.5'N, 94˚37.8'W, 155 m, 21 Jan 1975, P. Valdivia 1776 (XAL); Municipio Hidalgotitl&#225;n, 7 km going R&#237;o Alegre, 17˚12'N, 95˚04'W, 120 m, 26 Jan 1975, P. Valdivia 1888 (XAL); Municipio Hidalgotitl&#225;n, 4 km of Campamento Hermanos Cedillo to La Laguna, 17˚16'N, 94˚35'W, 160 m, 28 Jan 1975, P. Valdivia 1904 (XAL); Municipio Jes&#250;s Carranza, 10 km S of Hermanos Cedillo, road to R&#237;o Alegre, 17˚12'N, 94˚41'W, 150 m, 30 May 1974, M. V&#225;zquez 708 (XAL); Estaci&#243;n de Biolog&#237;a Tropical Los Tuxtlas, 19 mi (by road) NW of Catemaco, 18˚35'N, 95˚04'W, 400 ft [=122 m], 17 Aug 1976, G. Webster &amp; W. Armbruster 20888A (MEXU); Municipio Jes&#250;s Carranza, hills S of Pob. 2, (near 3 km S Of the junction of the earth trail La Laguna-Sarabia with the trail N to Pob. 2, 17˚12'N, 94˚39'W, 250 m, 14 Jul 1984, T Went &amp; A. Villalobos 4454 (CHAPA, MEXU); Municipio Hidalgotitl&#225;n, N of La Laguna camp, near and N Of the rubber plantation and the W of R&#237;o Cuevas, 17˚17'15''N, 94˚30'45''W, 125 m, 25 Sep 1980, T. Went et al. 2739 (CHAPA, MEXU).</p><p>Campyloneurum madrense differs from C. angustifolium by not pruinose (vs. pruinose) rhizome or scarcely so, linear (vs. lanceolate) and narrower (0.4 - 0.8 mm vs. 1.0 - 1.7 mm) rhizome scales with sparsely denticulate (vs. entire) margin, they dark gray to blackish (vs. yellowish-brown to grayish-brown), inside with narrow oblong (isodiametric) cells and present at lower (25 - 940 m vs. (700-) 1200 - 1600 (-2000) m) altitudinal distribution. Also the new species differs from C. gracile A. Rojas by thicker (2 - 5 mm vs. 1.5 - 2.5 mm) rhizome, longer (3 - 6 mm vs. 1.5 - 3.0 (-4.0) mm) rhizome scales with longer (2 - 5 times longer than wide vs. 1-2 times) cells and relative longer ((18-) 30 - 65 cm vs. (10-) 18 - 40 (-55) cm) fronds (see <xref ref-type="fig" rid="fig3">Figure 3</xref> and <xref ref-type="table" rid="table3">Table 3</xref>).</p><p>Studies on the type material of Campyloneurum angustifolium (Swartz s.n., holotype: S!; isotype: BM!), particularly on the isotype in BM is evident that this species has pruinose rhizome and yellowish-brown to brown rhizome scales, and also the material identified as C. ensifolium in Mexico not is significantly different, only in shorter and obtuse rhizome scales (present in the northernmost</p><fig id="fig3"  position="float"><label><xref ref-type="fig" rid="fig3">Figure 3</xref></label><caption><title> (a) (b). Type material of Campyloneurum madrense A. Rojas (D. &#193;lvarez et al. 6960, MEXU). (a) Habit; (b) Rhizome scales; (c) Blade detail</title></caption><graphic mimetype="image"   position="float"  xlink:type="simple"  xlink:href="http://html.scirp.org/file/19-2603096x4.png"/></fig><p>populations in Mexico), but this parameter is variable in the species gradually changing to South. However, Mickel &amp; Smith mentioned in the key and description of C. angustifolium from Mexico with oblong cells in rhizome scales and they twice or more longer than wide, characters more related with C. madrense, but also they mentioned populations of C. angustifolium in South of Mexico that are present at lower elevation and that have narrow rhizome scales with small cell lumen, and they associated this entity with C. angustipaleatum [<xref ref-type="bibr" rid="scirp.75089-ref7">7</xref>] , a species present in Costa Rica to Bolivia at high elevations.</p></sec></sec><sec id="s3_2"><title>3.2. Accepted Name and Lectotype Designed</title>Campyloneurum pittieri<p>(Christ) Ching, Sunyatsenia 5: 263. 1940 (<xref ref-type="fig" rid="fig4">Figure 4</xref>).</p><table-wrap id="table3" ><label><xref ref-type="table" rid="table3">Table 3</xref></label><caption><title> Comparison of morphological characteristics among three species of Campyloneurum: C. angustifolium, C. gracile and C. madrense</title></caption><table><tbody><thead><tr><th align="center" valign="middle" >Character</th><th align="center" valign="middle" >C. angustifolium</th><th align="center" valign="middle" >C. gracile</th><th align="center" valign="middle" >C. madrense</th></tr></thead><tr><td align="center" valign="middle" >Diameter of rhizome</td><td align="center" valign="middle" >2 - 5 mm</td><td align="center" valign="middle" >1.5 - 2.5 mm</td><td align="center" valign="middle" >2 - 5 mm</td></tr><tr><td align="center" valign="middle" >Pruinose rhizome</td><td align="center" valign="middle" >yes</td><td align="center" valign="middle" >not</td><td align="center" valign="middle" >not or scarcely so</td></tr><tr><td align="center" valign="middle" >Distance between fronds</td><td align="center" valign="middle" >approximate to 5 mm distant</td><td align="center" valign="middle" >approximate</td><td align="center" valign="middle" >approximate to 2 mm distant</td></tr><tr><td align="center" valign="middle" >Long of rhizome scales</td><td align="center" valign="middle" >3 - 5 mm</td><td align="center" valign="middle" >1.5 - 3.0 (-4.0) mm</td><td align="center" valign="middle" >3 - 6 mm</td></tr><tr><td align="center" valign="middle" >Broad of rhizome scales</td><td align="center" valign="middle" >1.0 - 1.7 mm</td><td align="center" valign="middle" >0.5 - 1.0 mm</td><td align="center" valign="middle" >0.4 - 0.8 mm</td></tr><tr><td align="center" valign="middle" >Margin of rhizome scales</td><td align="center" valign="middle" >entire</td><td align="center" valign="middle" >entire to denticulate</td><td align="center" valign="middle" >denticulate</td></tr><tr><td align="center" valign="middle" >Color of rhizome scales</td><td align="center" valign="middle" >yellowish-brown to grayish-brown</td><td align="center" valign="middle" >brown to dark brown</td><td align="center" valign="middle" >dark gray to blackish</td></tr><tr><td align="center" valign="middle" >Form of apical cells of rhizome scales</td><td align="center" valign="middle" >poligonal, elongate</td><td align="center" valign="middle" >poligonal, isodiametric</td><td align="center" valign="middle" >rectangular to oblong</td></tr><tr><td align="center" valign="middle" >Long of cell of rhizome scales</td><td align="center" valign="middle" >1.5 - 3.0 times longer than wide</td><td align="center" valign="middle" >1 - 2 times longer than wide</td><td align="center" valign="middle" >2 - 5 times longer than wide</td></tr><tr><td align="center" valign="middle" >Lumen color of rhizome scales</td><td align="center" valign="middle" >translucid</td><td align="center" valign="middle" >brown</td><td align="center" valign="middle" >gray</td></tr><tr><td align="center" valign="middle" >Frond size</td><td align="center" valign="middle" >20 - 45 (-75) cm</td><td align="center" valign="middle" >(10-) 18 - 40 (-55) cm</td><td align="center" valign="middle" >(18-) 30 - 65 cm</td></tr><tr><td align="center" valign="middle" >Diameter of stipe</td><td align="center" valign="middle" >1 - 2 mm</td><td align="center" valign="middle" >0.7 - 1.0 mm</td><td align="center" valign="middle" >1 - 1.5 (-2) mm</td></tr><tr><td align="center" valign="middle" >Broad of blade</td><td align="center" valign="middle" >0.4 - 1.5 (-2.0) cm</td><td align="center" valign="middle" >0.3 - 1.5 (-2.0) cm</td><td align="center" valign="middle" >(0.5-) 0.8 - 1.5 cm</td></tr><tr><td align="center" valign="middle" >Altitudinal distribution</td><td align="center" valign="middle" >(700-) 1200 - 1600 (-2000) m</td><td align="center" valign="middle" >0 - 900 (-1200) m</td><td align="center" valign="middle" >25 - 940 m</td></tr></tbody></table></table-wrap><fig id="fig4"  position="float"><label><xref ref-type="fig" rid="fig4">Figure 4</xref></label><caption><title> (a) (b). Representative specimen of Campyloneurum pittieri (Christ) Ching (A. Rodr&#237;guez &amp; L. Vargas 3152, CR). (a) Habit; (b) Rhizome scales; (c) Rhizome scales of Campyloneurum amphostenon (Kunze ex Klotzsch) F&#233;e (J. Bittner 109, CR)</title></caption><graphic mimetype="image"   position="float"  xlink:type="simple"  xlink:href="http://html.scirp.org/file/19-2603096x5.png"/></fig><p>Polypodium pittieri Christ, Prim. Fl. Costaric. 3(1): 16. 1901. Tipo: COSTA RICA. San Jos&#233;: El P&#225;ramo, E side of Cerro Buena Vista, 3000 m, Jan 1897, H. Pittier 10479 (lectotype designed here: CR!; isolectotypes: BM (photo, GH), BR, P!, US).</p><p>Description: Terrestrial; rhizome 2 - 5 mm in diameter, creeping, fronds 5 - 30 (-50) mm distant between them, pruinose; rhizome scales 2 - 5 &#215; 0.7 - 1.5 mm, ovate-lanceolate, clathrate, grayish-brown, sparse, peltate or with overlapping base, cells rounded to polygonal, lumen translucid; fronds 40 - 80 (-100) cm long, moderately decumbent; stipe 1/3 - 1/6 of the frond length, strawish to pale brown, glabrous, phyllopodia 2 - 5 mm long; blade 30 - 60 (-80) &#215; 2.5 - 3 (-5) cm, narrowly elliptic, long acuminate at apex, gradually reduced at base, coriaceous, lustrous; rachis glabrous, pale brown, pale as lamina, principally with immersed lateral veins or inconspicuous; sori rounded, in 3 - 7 rows between costa and margin.</p><p>Distribution: Known from Costa Rica and Panama at 700 - 1800 (-2000) m elevation.</p><p>Additional revised specimens: COSTA RICA. Cartago: Cerro Asunci&#243;n, ca. 0.5 km NE of the Interamerican Hwy (km 90 - 91), 6 Mar 1986, 3080 m, M. Kappelle 1414 (CR); Volc&#225;n Turrialba, outside the crater, 3200 - 3300 m, 16 Jun 1993, K. Mehltreter 432 (CR); El Guarco, Reserva Forestal Los Santos, Savegre basin, over Interamerican road, rute to Cerro de La Muerte, between km 78 and 79. 9˚37'05''N, 83˚49'0''W, 2960 m, 28 Jan 1998, A. Rodr&#237;guez et al. 2971 (INB, CR, MO); Volc&#225;n Turrialba, high part of the hill, 31 ene 1957, R. Rodr&#237;guez 98 (CR); Para&#237;so, Valle del Reventaz&#243;n, Cerros S&#225;kira, S&#225;bila and Zacatales, 9˚35'20''N, 83˚45'40''W, 3300 - 3420 m, 16 Jul 1996, A. Rojas &amp; M. Coto 2764 (INB, MO); Cerros Cueric&#237;, 2940 m, 9 Nov 1988, Y. Widmer 395 (CR). Cartago-San Jos&#233;: ca. 22 km SE of El Empalme, along Interamerican Hwy, 2500 - 2600 m, 9˚40'N, 83˚50'W, 27 Nov 1969, W. Burger &amp; R. Liesner 6501 (CR, F); just below of highest point on the Interamerican hwy, NW of La Asunci&#243;n, 9˚34'N, 83˚45'W, 3000 - 3200 m, 5 Feb 1982, W. Burger &amp; K. Barringer 11506 (CR, F); Cordillera de Talamanca, upper slopes, W ridge of Cerros Cueric&#237;, ca. 9˚34ʹN, 83˚40'W, 3160 m, 15 Sep 1983, G. Davidse 24661 (CR, MO); Cordillera de Talamanca, Cerros Cueric&#237;, Parque Nacional Chirrip&#243;, continental divide, 9˚35'N, 83˚38'30''W, 3200 m, 16 Sep 1983, G. Davidse 24813 (CR, MO); Villa Mills, 2 Aug 1961, D. Knight s.n. (CR). Heredia: Parque Nacional Braulio Carrillo, volc&#225;n Barva station, path and forest between park office and crater, 10˚07'20''N, 84˚06'00''W, 2700 - 2900 m, 13 Jul 1991, H. Gay et al. 1511 (CR); hills SW of Volc&#225;n Barba, 2700 - 2900 m, Apr 1971, L. G&#243;mez &amp; W. Wagner s.n. (CR); slopes and ridges above Laguna del Barva and Summit of Volc&#225;n Barva, 10˚08ʹN, 84˚06,5'W, 2840 - 2900 m, 27 Apr 1986, M. Grayum &amp; F. Quesada 7424 (CR, MO); Volc&#225;n Barva, trail to Laguna Copey, 2700 - 2800 m, 24 Jun 1993, K. Mehltreter 467 (CR); Barva, Parque Nacional Braulio Carrillo, T&#225;rcoles basin, Estaci&#243;n Barva, Sendero Secreto, 10˚08'00''N, 84˚07'15''W, 2600 m, 24 Mar 1998, A. Rodr&#237;guez &amp; L. Vargas 3152 (CR, INB, MO); Barva, Parque Nacional Braulio Carrillo, Sarapiqu&#237; basin, surroundings of the path that goes to the viewpoint, 10˚08'10''W, 84˚06'25''W, 1900 m, 2 Jun 1998, A. Rodr&#237;guez &amp; L. Vargas 3456 (CR, INB, MO); road from Cartago to San Isidro del General, (Pan American Hwy, Rte 2), km 87 - 88, ca. 1 km NW of Asunci&#243;n, 3200 m, 29 Jan 1986, A. Smith &amp; T. B&#233;liz 2022 (CR, UC); Aromal del Volc&#225;n Barba, 2700 m, 8 May 1927, M. Valerio 64 (CR). Lim&#243;n: Cordillera de Talamanca, Atlantic slope, K&#225;muk massif, paramo NE of the main K&#225;muk peak, 9˚16'N - 9˚17'N, 83˚00'W - 83˚02'W, 3000 - 3300 m, 17 - 18 Sep 1984, G. Davidse &amp; G. Herrera 29328 (CR, MO); cordillera de Talamanca, peak of Cerro K&#225;muk, 9˚16'30''N, 83˚02'W, 3350 - 3549 m, 25 Mar 1984, G. Davidse et al. 26019 (CR, MO); Chirrip&#243; National Park, N end of Loma Larga, ca. 3350 m, 15 Feb 1983, N. Garwood et al. 1263 (CR). Lim&#243;n-Puntarenas: Cordillera de Talamanca, between Cerro Kasir and Cerro Nai, continental divide, 9˚12'N - 9˚13'N, 83˚03'30''W - 83˚04'30''W, 2900 - 3050 m, 22 Mar 1984, G. Davidse et al. 25837 (CR, MO). San Jos&#233;: P&#233;rez Zeled&#243;n, Interamerican hwy (km 82) in the near of the Cerro de La Muerte border, 9˚34'20''N, 83˚45'30''W, 3000 m, 12 Jun 1991, J. Bittner 1097 (CR); along the trail from Cana&#225;n to Chirrip&#243;, via Los &#193;ngeles, above (N of) the R&#237;o Talari, 9˚30'N, 83˚32'W, 3100 - 3200 m, 24 Aug 1971, W. Burger 8323 (CR, F); along Interamerican Hwy, 9˚40'N, 83˚50'W, 2840 m, 23 Nov 1986, E. Hennipman et al. 7131 (CR, U); Cerro Las Vueltas, between Cruces and Alto Indio, 3020 m, 11 Mar 1986, M. Kappelle 1622 (CR); Parque Nacional Chirrip&#243;, Monte Sin F&#233;, 3250 m, 20 Sep 1988, M. Kappelle 2674 (CR); Parque Nacional Chirrip&#243;, Palmito Morado, 2910 m, 20 Jan 1989, M. Kappelle &amp; M. Monge 4902 (CR); San Gerardo de Dota, 2800 m, Jun-Jul 1989, H. Van Velzen &amp; T. Genze 62 (CR).</p><p>PANAM&#193;. Bocas del Toro: Cordillera de Talamanca, 2 airline km NW of the main peak of cerro F&#225;brega, along the NW ridge of the massif, 9˚08'N, 82˚53'W, 3150 - 3200 m, 7 - 8 Mar 1984, G. Davidse et al. 25333 (CR, MEXU, MO); between Itamut &amp; Bine peaks, Fabrega massiff, 3200 m, 5 - 9 Mar 1984, L. G&#243;mez et al. 22547 (CR); 1 - 2 km SWW of Itamut camp, 3175 m, 6 - 7 Mar 1984, L. G&#243;mez et al. 22615 (CR).</p><p>Campyloneurum pittieri differs from C. amphostenon because has not pruinose or scarcely pruinose (vs. pruinose); rhizome scales longer (4 - 8 mm vs. 2 - 5 mm), ovate with a long acuminate tip (vs. lanceolate with acute tip), they are with brown to dark brown color (vs. yellowish-brown to grayish-brown) and with the apical cells 1 - 2 times longer than wide (vs. 1.5 - 3 times); fronds relatively longer ((35-) 45 - 60 (-80) cm vs. 30 - 55 (-65) cm); stipe longer (8 - 25 cm vs. 4 - 10 (-20) cm); blade relatively broader ((1.3-) 2 - 5 cm vs. 1 - 3.5 cm); veins prominently marked (vs. not marked or scarcely so) and distributed to relative higher elevation ((1900-) 2700 - 3400 (-3550) m vs. (1800-) 2100 - 2900 (-3200) m) (see <xref ref-type="fig" rid="fig4">Figure 4</xref> and <xref ref-type="table" rid="table2">Table 2</xref>).</p><p>Detailed studies on the type of Campyloneurum amphostenon (Moritz 120b, B!, BK!, K!) demonstrated that this material coincides with the second entity proposed by Rojas (2005), while the first corresponds to C. pittieri (Christ) Ching (H. Pittier 10479, CR!), so the recognition and validation of this species are necessary.</p></sec></sec><sec id="s4"><title>Acknowledgements</title><p>This work was made possible thanks to Universidad Nacional de Costa Rica (UNA) and Sistema Nacional de &#193;reas de Conservaci&#243;n (SINAC). Thanks also to Herbario Nacional de Costa Rica (CR) and Herbario Nacional de M&#233;xico (MEXU) for providing specimen loans and the space for analyze the specimens. For the good attention and facilitates of the curators in the Mexican herbaria CHAPA, MEXU and XAL. Thanks to Pablo Mu&#241;oz Cambronero, Jasm&#237;n Mej&#237;a Vargas and M&#243;nica Guzm&#225;n Morales for the excellent drawings. To the anonymous reviewers for their help; and to all who, in one way or another, made this work possible.</p></sec><sec id="s5"><title>Cite this paper</title><p>Rojas-Alvarado, A.F. (2017) Novelties in Campyloneurum (Polypodiaceae) from Mesoamerica. 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