This article was downloaded by: [Alessandro Catenazzi] On: 02 August 2012, At: 07:51 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK Diatom Research Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/tdia20 Frankophila wayqechae sp. nov., a new aerophilic diatom species from the Peruvian Andes, South America Paula C. Furey a b , Shigeki Mayama c , Rex L. Lowe d & Alessandro Catenazzi b e a Department of Biology, St. Catherine University, St. Paul, MN, USA b Department of Integrative Biology, University of California, Berkeley, CA, USA c Department of Biology, Tokyo Gakugei University, Tokyo, Japan d Department of Biological Sciences, Bowling Green State University, Bowling Green, OH, USA e Department of Biology, San Francisco State University, San Francisco, CA, USA Version of record first published: 24 Jul 2012 To cite this article: Paula C. Furey, Shigeki Mayama, Rex L. Lowe & Alessandro Catenazzi (2012): Frankophila wayqechae sp. nov., a new aerophilic diatom species from the Peruvian Andes, South America, Diatom Research, 27:3, 165-175 To link to this article: http://dx.doi.org/10.1080/0269249X.2012.704884 PLEASE SCROLL DOWN FOR ARTICLE Full terms and conditions of use: http://www.tandfonline.com/page/terms-and-conditions This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae, and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand, or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material.
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This article was downloaded by: [Alessandro Catenazzi]On: 02 August 2012, At: 07:51Publisher: Taylor & FrancisInforma Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House,37-41 Mortimer Street, London W1T 3JH, UK
Diatom ResearchPublication details, including instructions for authors and subscription information:http://www.tandfonline.com/loi/tdia20
Frankophila wayqechae sp. nov., a new aerophilicdiatom species from the Peruvian Andes, SouthAmericaPaula C. Furey a b , Shigeki Mayama c , Rex L. Lowe d & Alessandro Catenazzi b ea Department of Biology, St. Catherine University, St. Paul, MN, USAb Department of Integrative Biology, University of California, Berkeley, CA, USAc Department of Biology, Tokyo Gakugei University, Tokyo, Japand Department of Biological Sciences, Bowling Green State University, Bowling Green, OH,USAe Department of Biology, San Francisco State University, San Francisco, CA, USA
Version of record first published: 24 Jul 2012
To cite this article: Paula C. Furey, Shigeki Mayama, Rex L. Lowe & Alessandro Catenazzi (2012): Frankophila wayqechae sp.nov., a new aerophilic diatom species from the Peruvian Andes, South America, Diatom Research, 27:3, 165-175
To link to this article: http://dx.doi.org/10.1080/0269249X.2012.704884
PLEASE SCROLL DOWN FOR ARTICLE
Full terms and conditions of use: http://www.tandfonline.com/page/terms-and-conditions
This article may be used for research, teaching, and private study purposes. Any substantial or systematicreproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form toanyone is expressly forbidden.
The publisher does not give any warranty express or implied or make any representation that the contentswill be complete or accurate or up to date. The accuracy of any instructions, formulae, and drug doses shouldbe independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims,proceedings, demand, or costs or damages whatsoever or howsoever caused arising directly or indirectly inconnection with or arising out of the use of this material.
Diatom ResearchVol. 27, No. 3, September 2012, 165–175
Frankophila wayqechae sp. nov., a new aerophilic diatom species from the Peruvian Andes,South America
PAULA C. FUREY1,2∗, SHIGEKI MAYAMA3, REX L. LOWE4 & ALESSANDRO CATENAZZI2,5
1Department of Biology, St. Catherine University, St. Paul, MN, USA2Department of Integrative Biology, University of California, Berkeley, CA, USA3Department of Biology, Tokyo Gakugei University, Tokyo, Japan4Department of Biological Sciences, Bowling Green State University, Bowling Green, OH, USA5Department of Biology, San Francisco State University, San Francisco, CA, USA
The genus Frankophila Lange-Bertalot is characterized as a small chain-forming diatom with frustules held together by interdigitatingmarginal spines. Both valves of the species in this genus possess a short raphe at the distal ends. Five species of Frankophila have beenrecognized to date with most being described from subaerial habitats. Exploration of moist, vertical, rock walls in the Peruvian Andes ledto the discovery of a new Frankophila species, F. wayqechae Furey, Mayama, Lowe & Catenazzi, sp. nov., which is described from a highaltitude (3506 m above sea level), moist, vertical wet wall located in the transition zone between the grasslands and cloud forests growingon the eastern slope of the Andes in Manu National Park. Light and scanning electron micrographs of internal and external morphologyand ultrastructure are presented along with basic ecological information. This new taxon is compared with species in the genus HygropetraKrammer & Lange-Bertalot, primarily H. gelasina Mayama & Idei, and phylogenetic relationships inferred from valve and chloroplastfeatures are discussed. The triseriate striae and depressions in the valve axial area of F. wayqechae differentiate it from other currentlydescribed Frankophila species. The velum at the inner opening of the areolae and triseriate striae of F. wayqechae are similar to the genusHygropetra which suggests a close relationship between these taxa. Frankophila wayqechae has a single chloroplast per cell that expandsin the large area beneath one valve without lobes along a sternum. The nature of the chloroplast, combined with the pattern of areolationin the striae and internal areola occlusion by a hymen also suggests F. wayqechae may be a close relative of Planothidium Round &Bukhtiyarova. Frankophila wayqechae appears to be restricted to high elevation areas in the Peruvian Andes, and is likely an endemictaxon, emphasizing the importance of biodiversity hotspot studies for future species conservation.
Keywords: Frankophila, Hygropetra, Andes, Peru, wet walls, subaerial habitats
IntroductionLange-Bertalot (1997) described the diatom genusFrankophila based on samples from Chile and the USA.Five species of Frankophila have been recognized todate with most being described from subaerial habitats:F. similioides Lange-Bertalot & Rumrich, F. loetschertii(Lange-Bertalot) Lange-Bertalot, F. maillardii (Le Cohu)Lange-Bertalot, F. horstii Rumrich & Rumrich andF. biggsii Lowe, Morales & Kilroy. The genus Frankophilais characterized as a small chain-forming diatom with frus-tules held together by interdigitating marginal spines, whichhave been to date located on virgae. Both valves in thisgenus possess a short raphe at its distal ends.
A new species of Frankophila is described fromhigh altitude, subaerial habitats in the Peruvian Andes,including detailed light (LM) and scanning electron(SEM) micrographs of the internal and external mor-phology and ultrastructure, along with general ecologicalinformation. Morphological features and taxonomy of this
new species are compared with other Frankophila speciesand with species in the genus Hygropetra Krammer &Lange-Bertalot, primarily H. gelasina Mayama & Idei. Thephylogeny inferred from valve and chloroplast features isdiscussed.
Materials and methodsDiatom samples were collected from 23 moist subaerialhabitats (wet walls) and two streams along an elevationgradient from 1615 to 3506 m above sea level (a.s.l.) inthe montane cloud forest, montane scrub and high-Andeangrasslands in Manu National Park, Department of Cusco,southeastern Peru in February 2009 and January 2010(Table 1). Two composite samples were collected frommost of the wet wall sites, one by scraping material fromrock surfaces with a spoon, and one by squeezing mate-rial from mosses and hepatics into a plastic bag. Wet rockwalls were designated wet if water was visibly running overthe rock face and moist if not. Lotic composite samples
Notes: a.s.l., above sea level; Scr., scraping from wet walls; Br., bryophyte squeeze; Bold text, type locality of Frankophila wayqechaesp. nov. Moisture level as defined in Materials and methods.
were collected by scraping periphyton from three to fourrepresentative cobbles in the streams. All samples weresplit in two with one set kept moist and the other air-driedbefore shipping back to USA and Japan for microscopicanalysis.
The type locality, a wet rock wall at an elevation of3506 m, is located along the Paucartambo–Shintuya roadnear the mountain pass of Abra Acjanaco. This pass crossesthe ridge that separates the wet eastern slopes of theCordillera de Paucartambo from the drier inter-Andeanvalley of Paucartambo. The type locality is found in thetransition zone between the high Andean grasslands (puna)and cloud forests growing on the eastern slope of the Andes,in the headwaters of the Kosñipata River in Kosñipata Val-ley, on the eastern side of the Cordillera de Paucartambo.Here the treeline is generally between 3200 and 3400 m butreaches 3550 m in protected areas and along streams wherewet walls are commonly found. The wet wall is exposed tothe east and receives water run-off from the nearby grass-land and elfin forest throughout the year, although it isreduced during the dry season. Vegetation of the upperKosñipata Valley includes trees of the genera Alnus Miller,Clethra Linnaeus, Weinmannia Linnaeus, Clusia Linnaeus,Symplocos Jacquin, bamboos (Chusquea Kunth), arborealferns (Cyathea Smith), terrestrial and epiphytic bromeliads(Puya Molina, Pitcairnia L’Héritier, Tillandsia Linnaeus),ericaceous shrubs, and herbs, vines and epiphytes of the
genera Begonia Linnaeus, Bomarea Mirbel, CalceolariaLinnaeus, Oxalis Linnaeus and Peperomia Ruiz & Pavon(Cano et al. 1995). The bedrock at elevations > 2000 m isprimarily slate or mudstone, at 1800 to ca. 2000 m it isigneous rock or basalt (no samples were collected fromthese elevations) and from 1500 to 1800 m it is mostlygranitic substrate (Table 1; K. Clark, unpubl., Universityof Oxford).
Samples were cleaned in sulfuric acid with potassiumdichromate or by boiling nitric acid in a 1:1 sample toacid ratio until the volume returned to the original vol-ume of 25 mL. The samples were rinsed with distilled waterand allowed to settle for a minimum of 8 h before decant-ing and fresh rinsing. This rinsing process was repeatedca. 10 times until the pH of the suspension was neutral.The cleaned material was then concentrated, air-dried ontocoverslips and processed for either light (LM) or scanningelectron microscope (SEM) analysis. Strewn diatom slidesmade using Naphrax� mounting medium were examinedat 1000× under an Olympus BX51 Photomicroscope withhigh resolution Nomarski (DIC: differential interferencecontrast) optics (Olympus America, Melville, NY, USA).The relative abundance of diatoms was determined bycounting a minimum of 600 valves at 1000× in discretefields of view. Images were captured with a monochro-matic camera (Spot™, Diagnostic Instruments, Inc., USA)attached to the microscope. Size variability (length, width,
Chloroplasts were examined at 1000× from aggrega-tions of putative living specimens of Frankophila from2010. Cells containing chloroplasts were then confirmedto belong to Frankophila by drying the aggregations ona microscope slide and examining them under SEM afterprocessing. To clean the targeted diatoms, the coverslip wascarefully removed and drops of diluted sodium hypochloritesolution were added to the Frankophila aggregations. Oneminute later, using a micropipette under an inverted micro-scope the solution was removed and rinsed several timeswith distilled water. These specimens were transferred to analuminum stub and coated with 5 nm of osmium tetroxideusing OPC-60A Osmium Plasma Coater (Filgen, Nagoya,Japan) for SEM observations.
Descriptio. Frustulae rectangulares aspectu cinguli for-mantes filamenta connexa spinis marginalibus interdig-itatis. Valvae ellipticae apicibus late rotundatis. Longi-tudo 4.5–8.0 μm, latitudo 3.0–4.6 μm, altitudo limbi 1.3–3.0 μm. Area axalis lata, depressionibus vadosis irregu-lariter positis in facie externa. Rami raphis <1 μm longi,positi in terminatione distali valvae. Striae radiatae omnino,11–12 in 10 μm, constantes ex seriebus tribus areolarum,extensis in limbo valvae. Longitudo striae 45–47% latitu-dinis valvae. Interne, areolae tectae hymenibus perforatis.Spinae solidae-granulosae (non subtiliter siliceae, mag-nitudo granularum 5–10 nm), dichotome ramosae, posi-tae in virgis. Valvocopula perforate rimis elongatis propemarginem limbi.
Description. Frustules rectangular in girdle viewforming filaments held together by interdigitating marginalspines. Valves elliptical with broadly rounded apices invalve view. Length 4.5–8.0 μm, width 3.0–4.6 μm, per-valvar axis 1.3–3.0 μm, stria density 11–12 in 10 μm.
Combined stria length on average 45–47% of the width ofthe valve face. Valves with a broad axial area with shallowdepressions arranged in various patterns on the external sur-face. Striae composed of three rows of areolae which extendonto the valve mantle. There is a hymen at the inner openingof each areola which has fine perforations (ca. 5 nm). Solidspines dichotomously branched, located on the virgae. Veryshort raphe slits <1 μm long, on distal end of valve. Valvo-copula open, with slits near the mantle edge accompaniedby three narrow, plain pleurae.
Holotype. Circled specimen on slide ANSP GC26816deposited in the Diatom Herbarium of the Academy ofNatural Sciences (ANSP), Philadelphia, USA. Figure 9 ishere designated as the holotype. Type material depositedat ANSP (ANSP GCM10307), the Diatom Herbariumof the California Academy of Sciences, San Francisco,USA (CAS 627413), the National Algal Collectionat the Canadian Museum of Nature, Ottawa, Canada(CANA 85058) and the Department of Botany, NationalMuseum of Nature and Science, Tokyo, Japan (TNS-AL-56980).
Isotypes. Circled specimens on slides deposited in theDiatom Herbarium of the California Academy of Sci-ences, San Francisco, USA (CAS 223025, Fig. 8), theNational Algal Collection at the Canadian Museum ofNature, Ottawa, Canada (CANA 85058, Fig. 12) and theDepartment of Botany, National Museum of Nature andScience, Tokyo, Japan (TNS-AL-56980).
Type locality. In mucilage on a vertical, wet rock wallat an elevation of 3506 m a.s.l. along the Paucartambo–Shintuya road near the Abra Acjanaco mountain pass, ManuNational Park, Department of Cusco, southeastern Peru(13.2016◦S, 71.6167◦W) (Table 1). Collected by A. Cate-nazzi, 13 February 2009 as sample Peru33 – PC Furey(personal collection).
Etymology. The specific epithet refers to the Quechuaword for friend for the Wayqecha Biological Station, ownedand operated by the Asociación para la Conservación de laCuenca Amazónica.
Observations. The frustules are rectangular in girdleview and form filaments held together by interdigitatingmarginal spines (Figs 1–2, 20–23, arrows, 26). The valvesare small and elliptical with broadly rounded apices in valveview (Figs 1–20, 24–25). The valves range in length from4.5 to 8.0 μm and in width from 3.0 to 4.6 μm. The perval-var axis varies from 1.3 to 3.0 μm, and the stria density is11–12 in 10 μm. The combined stria length on average is45–47% of the width of the valve face. The valves have abroad axial area that has shallow depressions arranged invarious patterns on the external surface (Figs 4–20, 24–25,black arrows). The striae are composed of three rows ofareolae which extend onto the valve mantle (Figs 24–27).There is a hymen at the inner opening of each areola whichhas fine perforations ca. 5 nm (Figs 28, 30–31). The valveshave solid spines that are dichotomously branched and arelocated on the virgae (Figs 24, 26–27, 29–30). Very short
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Figs 1–23. Type material of Frankophila wayqechae from Manu National Park, southeastern Peru. Figs 1–3. Live cells with chloroplastsin valve and girdle views. Figs 4–19. Variation of the valve surface in valve views. Fig. 8. Isotype specimen, CAS 223025. Fig. 9. Holotypespecimen, ANSP GC26816. Fig. 12. Isotype specimen, CANA 85058. Fig. 20. Valve and frustule in girdle view. Figs 21–23. Frustulesforming ribbon-shaped colonies in girdle view, with some spines visible (arrows). Scale bars = 10 μm (Figs 1–3), 5 μm (Figs 4–23).
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Figs 24–31. Type material of Frankophila wayqechae from Manu National Park, southeastern Peru, external (Figs 24–30) and internalview (Fig. 31). Figs 24–25. Whole valves showing striae, spine position on virgae, raphe slits (white arrows) and shallow surface depressions(black arrows). Fig. 26. Paired frustules in girdle view linked by interdigitating marginal spines, showing girdle bands, mantle striae anddichotomously branched spines (arrow). Fig. 27. Valve margin showing a valvocopula (VC) and three pleurae (P1–P3). Fig. 28. Hymenateareolae with fine perforations (ca. 5 nm in diameter). Figs 29–30. Broken solid spines. Fig. 31. Part of valve showing striae and short raphefissure (arrow). Scale bars = 2 μm (Fig. 26), 1 μm (Figs 24–25, 27, 31), 0.2 μm (Figs 28–30).
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raphe slits (<1 μm long) are located on the distal end of thevalve (Figs 24–25, 31, white arrows). The valvocopula isopen, with slits near the mantle edge (Figs 24, 26–27) thatare accompanied by three narrow, plain pleurae (Fig. 27).There is a single chloroplast per cell that lies in a more or lesscurved plate beneath one valve, folding down appressed toone side of the girdle and expanding a bit to the other valve(Figs 1–3); no pyrenoids were detected.
Habitat and distribution. Frankophila wayqechaeformed aggregates (solitary or colonial, Figs 21–23) on therocks or mucilage covering mosses and hepatics, ca. 10–20 cm from flowing water at the type locality. Based on 600valves counted from each sample from all of the locations,F. wayqechae was only found at the type locality. The rel-ative abundance of F. wayqechae cells at the type locality,determined from four samples collected in 2009 and 2010was 80–88% in three samples and 25% in one sample.
DiscussionOther known subaerial Frankophila species, such asF. biggsii are restricted to particular rock types (Lowe et al.2006). Rock walls in this study at elevations >2000 m a.s.l.such as the type locality, are primarily composed of slateor mudstone. However, F. wayqechae was absent from the16 other wet wall sites and two streams collected at thesehigher elevations. Frankophila wayqechae was also absentfrom six wet walls collected at altitudes <2000 m a.s.l.,which are composed of other mineral types (e.g., granite).Without further study, it is unclear whether F. wayqechaeis restricted to high altitude sites (i.e., at altitudes near orgreater than 3000 m a.s.l.) or if it is restricted to a particularrock type or other unique suite of habitat characteristics.
Comparison of F. wayqechae with other FrankophilaspeciesFrankophila wayqechae shares some habitat and mor-phological similarities with other currently describedFrankophila taxa, but can readily be differentiated (Table 2).No other currently described Frankophila species has trise-riate striae and depressions in the valve sternum. Additionaldifferences and similarities are summarized in Table 2 andhighlighted below.
Frankophila similioides and F. wayqechae both have anelliptical valve with broad rounded apices, but F. wayqechaehas a broader sternum than F. similioides (Lange-Bertalot1997, Rumrich et al. 2000, Metzeltin & García-Rodríguez2003, Metzeltin et al. 2005). The valves of F. wayqechaehave a slightly shorter length and narrower width, and ashorter raphe than F. similioides. The striae of F. wayqechaeare less dense than those of F. similioides. Both possess slit-like perforations on the pars exterior and near the valvemantle edge. Frankophila wayqechae and F. similioidesare both subaerial diatoms from South America. However,
F. similioides has also been observed in non-subaerial habi-tats in South America such as the hot springs of La Calera inthe Peruvian Andes (Cocquyt & Van de Vijver 2007) andhigh elevation streams in Bolivia (2450 m a.s.l., Moraleset al. 2009; 4056 m a.s.l., McClintic et al. 2003, Moraleset al. 2007) and Argentina (Rio Cortadera at 4000 m a.s.l.and Rio Mirihuaca at 3420 m a.s.l., Maidana & Seeligmann2006).
Frankophila loetschertii valves are longer and broaderthan the valves of F. wayqechae (Lange-Bertalot 1997).The striae of F. loetschertii extend farther onto the valveface and contain two, occasionally three, rows of areolaein contrast with the shorter striae of F. wayqechae, whichhave three rows of areolae. The virgae of F. wayqechae areraised to a lesser degree than the conspicuously raised vir-gae of F. loetschertii; however, both taxa have bifurcatingspines located on the virgae. Little additional informationis available on the distribution of F. loetschertii.
Frankophila maillardii valves have wide areolae andwell-developed striae which result in a narrow central ster-num in contrast to the round areolae and shorter striae ofF. wayqechae, which yield a broader axial area (Lange-Bertalot & Le Cohu 1985, Le Cohu 1999). Similar to thewet subaerial habitat of F. wayqechae, F. maillardii has beenreported in low numbers from seeps and springs from thesub-Antarctic Prince Edward Islands (Van de Vijver et al.2008). In contrast, F. maillardii has also been found as adominant taxon in streams in sub-Antarctic Crozet Islands(Van de Vijver & Beyens 1999).
Frankophila horstii has a cruciate valve outline with alanceolate to rhombic central sternum that differs markedlyfrom the broadly elliptical valve outline and broadly ellip-tical axial area of F. wayqechae (Rumrich et al. 2000). Thevalves of F. wayqechae are smaller than those of F. horstii.The striae of F. wayqechae contain three rows of areolaein contrast with the double rows of areolae in F. horstii.Additional information on the distribution of F. horstii wasnot found.
Frankophila biggsii valves are longer in length and elon-gate elliptical in valve outline compared to the shorter,broadly elliptical valves of F. wayqechae (Lowe et al. 2006).The striae of F. biggsii contain double rows of areolae incontrast with the three rows of areolae in F. wayqechae.Both taxa are described from subaerial wet walls, and havenot been reported from other habitat types or locations.
Relationship of F. wayqechae to HygropetraThe close similarity of the genus Hygropetra to Frankophilahas already been pointed out in the establishment of thisgenus; however, the shorter raphe branches of Frankophila
Table 2. Comparisons of the ecology, biometric data and morphological features between the new species Frankophila wayqechae and related taxa in the genera Frankophila andHygropetra.
F. wayqechae sp. nov. F. biggsii F. horstiia F. loetschertii F. maillardiib F. similioides H. balfouriana H. elongata H. gelasina
Type locality Manu NationalPark,PeruvianAndes
Fox River,Punakaiki,New Zealand
Nacional Park LasCajas, Ecuador
Firehole River,YellowstoneNationalPark, USA
Lac desKorrigans,KerguelenIslands
ChileanPatagonia
FreshwaterScotland,UKc
Firehole River,Wyoming,USA
RyugaeshiFalls, Japan
Habitat Wet rock wall Wet rock wall ‘habitat similar toFragilaria leptostauron’d
Running water Lakes Small, shallowpool,subaerial
Epiphytic onbryophytes,epipelice
Running water Epiphytic onbryophytesnearwaterfalls
n.d. n.d. Distal rapheends a shortdistancefrom eachapex
n.d. n.d.
Notes: n.d., no data. a Determined from SEM micrographs. b From Lowe et al. (2006). c Based on Cleve (1895). d As described in Rumrich et al. (2000); Fragilaria leptostauron(Ehrenberg) Hustedt is a synonym of Staurosirella leptostauron (Ehrenberg) Williams & Round. e Based on Mayama & Idei (2009). f Morphometry from Kociolek (2011).
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were still distinct criteria for separating the two genera(Krammer 2000). The discovery of H. gelasina throwsdoubts on this generic diagnosis (Mayama & Idei 2009),because that species has rudimentary short raphe branchesdifferent from the other Hygropetra species (Table 2).Mayama & Idei (2009) discussed the similarity of thehexagonal pattern of the sub-pores in the striae, which werepartly borne in some Frankophila species, and suggested aclose relationship between the two genera. In F. wayqechae,the hymen at the inner opening of the areolae, the areola pat-tern in the striae and the depressions on the valve face arenew information to enhance the relationship between thesetwo genera (Table 2).
However, the presence of linking spines is a charac-teristic distinguishing F. wayqechae from H. gelasina inwhich spines are absent. Mayama & Idei (2009) discussedthe stability of the linking spines as a generic criterion withthe evidence of interdigitating spines generated by EunotiaEhrenberg and Pinnularia Ehrenberg species, suggestingthat Frankophila is not strictly defined by the posses-sion of linking spines. Their statement is also supportedby the presence of other spine-bearing Pinnularia species(Van de Vijver et al. 2004, 2009). Information about thechloroplast of Hygropetra is still lacking. Explorations ofthe live cells as well as molecular analyses could pro-vide additional information on the potential phylogeneticrelationships between these taxa.
Valve, chloroplast and habitat featuresFrankophila was originally considered to belong in thefamily Fragilariaceae Greville because of prominent link-ing spines forming a colony despite the presence of simplelongitudinal slits at both valve ends and facultative spinediminution or loss (Lange-Bertalot 1997). However, thecombination of the raphe slits and loculate areolae observedin F. biggsii required the transfer of this genus to a raphidorder (Lowe et al. 2006), although the detail of areola occlu-sions was not mentioned. As pointed out by Mayama & Idei(2009), an ambiguous image of a velum-like structure wasshown in a specimen of Frankophila (Lange-Bertalot &Le Cohu 1985, fig. 32), but reliable information to confirmthis areola occlusion has not been available. The hymenatepore occlusion observed in the present study assures theinclusion of this genus as a member of the raphid diatoms,because the hymen is one of the characters distributed inmany raphid genera (Cox 2004).
The number and configuration of the chloroplasts,along with the valve morphology observed in Frankophila,confine its phylogenetic position. The single chloroplastobserved in F. wayqechae confirms its exclusion fromthe Fragilariaceae, because chain-forming araphid diatomsusually have two plate-like chloroplasts, e.g., Fragilaria,Staurosirella Williams & Round, Staurosira Ehrenbergand Punctastriata Williams & Round (Round et al. 1990,Cox 1996). Among raphid taxa with a single chloroplast
per cell, its configuration is categorized into several types(Cox 1996), which seem to correspond well with currentlyavailable phylogenetic clades (Bruder & Medlin 2008,Bruder et al. 2008). The configuration of the chloroplastin F. wayqechae has similarity to those of the monoraphidgenera Achnanthidium Kützing, Cocconeis Ehrenberg andPlanothidium (Cox 1996), in which the chloroplast expandsin the large area beneath one valve without lobes along asternum. In biraphid diatoms, chloroplasts are absent fromthe position beneath the raphe or are reduced due to lobes.The formation of ribbon colonies might have led to the lossof motility in Frankophila as indicated by the remainingrudimentary raphe and the chloroplast lying under one valveface. Therefore, the similar configuration of the chloro-plasts between Frankophila and the monoraphid diatomsmay be an evolutionary convergence, but further research isrequired to confirm this hypothesis. However, the hexagonalareolation in the striae and the internal areola occlusion byhymenes are shared features that are part of the overall char-acterization of the monoraphid genus Planothidium (Round& Bukhtiyarova 1996, Kobayasi et al. 2006). Similar shar-ing of morphological characteristics also occurs betweenPlanothidium and Hygropetra, which was previously sug-gested to be a close relative to Frankophila (Mayama &Idei 2009).
Similar to F. wayqechae, and also F. similioides andF. biggsii, both H. balfouriana (Grunow ex Cleve)Krammer & Lange-Bertalot and H. gelasina have beenreported from subaerial environments (Mayama & Idei2009). However, Frankophila taxa have generally beenreported from locations outside North America (see discus-sion above), whereas H. balfouriana is broadly distributedacross a variety of habitats, including lakes and rivers inNorth America (e.g., Kociolek 2011) and around the world(e.g., Bouchard et al. 2004, Rott et al. 2006). Because thefeatures used to distinguish Hygropetra species are notreadily visible under the LM (Morales 2003), reports ofH. gelasina and H. elongata Krammer & Lange-Bertalotare less common so comparison with Frankophila speciescannot yet readily be made.
Tropical mountain tops in Peru and throughout theAndes are areas of high species richness and endemismfor many plant and animal groups (e.g., Cano et al. 1995,Patterson et al. 1998, Young & León 2000), includingdiatoms (Manguin 1964, Rumrich et al. 2000). The biodi-versity, ecosystem integrity and health of both terrestrial andaquatic ecosystems in this biodiversity hotspot are threat-ened by the intensification of human land use, the spreadof invasive species (Brown 2003, Catenazzi et al. 2011)and from predicted changes in moisture and temperaturewith climate change (Vuille et al. 2003, Urrutia & Vuille2009). The new diatom species F. wayqechae appears to berestricted to high elevation areas in the Peruvian Andes andis likely an endemic taxon, emphasizing the importance ofbiodiversity hotspot studies for conservation. Furthermore,taxa like F. wayqechae, found on habitats dependent on air
moisture and groundwater flow such as wet walls, may beuseful bioindicators of shifts in moisture and temperaturepatterns in high elevation areas in Peru predicted to occurwith climate change (Urrutia & Vuille 2009). Given the highbiodiversity of this area, the authors predict the discoveryof more new diatom taxa from the subaerial samples in thisstudy and encourage more biodiversity studies in the region.
AcknowledgementsA special thank you to Sandra Almeyda and Juan Carlos Jahuanchifor assistance with collecting samples from Peru. We acknowledgethe Manu National Park, Peru for granting research permits and theWayqecha Biological Station for logistic support. Fieldwork wassupported with grants from the Amazon Conservation Associa-tion and the Rufford Small Grants Foundation. AC was supportedby a Postdoctoral Fellowship from the Swiss National ScienceFoundation (#116305). PCF received postdoctoral fellowship sup-port in part by the National Center for Earth-surface Dynamics(NCED) and a National Science Foundation grant awarded to JillWelter [NSF-DEB 0950016] of St. Catherine University. Thankyou to Bowling Green State University, Department of BiologicalSciences for access to and use of their scanning electron micro-scope. Thank you to Bart Van de Vijver of the National BotanicGarden of Belgium for the Latin translation.
ReferencesAnonymous. 1975. Proposal for standardization of diatom
terminology and diagnoses. Beihefte zur Nova Hedwigia 53:323–354.
Bouchard G., Gajewski K. & Hamilton P.B. 2004. Freshwaterdiatom biogeography in the Canadian Arctic Archipelago.Journal of Biogeography 31: 1955–1973.
Brown L. 2003. Plan B: rescuing a planet under stress and acivilization in trouble. W.W. Norton and Co., New York.285 pp.
Bruder K. & Medlin L.K. 2008. Morphological and molecularinvestigations of naviculoid diatoms. II. Selected genera andfamilies. Diatom Research 23: 283–329.
Bruder K., Sato S. & Medlin L.K. 2008. Morphological andmolecular investigations of naviculoid diatoms. IV. Pinnu-laria vs. Caloneis. Diatom 24: 8–24.
Cano A., Young K.R., León B. & Foster R.B. 1995. Composi-tion and diversity of flowering plants in the upper montaneforest of Manu National Park, southern Peru. In: Biodiversityand conservation of neotropical montane forests (Ed. by S.P.Churchill, H. Balslev, E. Forero & L.L. Luteyn), pp. 271–280.New York Botanical Garden, New York.
Catenazzi A., Lehr E., Rodriguez L.O. & Vredenburg V.T.2011. Batrachochytrium dendrobatidis and the collapse ofanuran species richness and abundance in the upper ManuNational Park, southeastern Peru. Conservation Biology 25:382–391.
Cleve P.T. 1895. Synopsis of the naviculoid diatoms. Part II.Kongliga Svenska-Vetenskaps Akademiens Handlingar 27:1–219.
Cocquyt C. & Van de Vijver B. 2007. La Calera: diatom com-position of a Peruvian hot spring in the Colca canyon. In:Proceedings of the 1st Central European diatom meeting (Ed.
by W.-H. Kusber & R. Jahn), pp. 31–33. Botanic Garden andBotanical Museum Berlin-Dahlem, Freie Universität Berlin.
Cox E.J. 1996. Identification of freshwater diatoms from livematerial. Chapman & Hall, London. 158 pp.
Cox E.J. 2004. Pore occlusions in raphid diatoms – a reassessmentof their structure and terminology, with particular referenceto members of the Cymbellales. Diatom 20: 33–46.
Cox E.J. & Ross R. 1980. The striae of pennate diatoms. In: Pro-ceedings of the sixth symposium on recent and fossil diatoms(Ed. by R. Ross), pp. 267–278. Otto Koeltz, Koenigstein.
Kobayasi H., Idei M., Mayama S., Nagumo T. & Osada K. 2006.H. Kobayasi’s atlas of Japanese diatoms based on electronmicroscopy. Volume 1. Uchida Rokakuho Publishing, Tokyo.533 pp.
Kociolek P. 2011. Hygropetra balfouriana. In: Diatoms ofthe United States. http://westerndiatoms.colorado.edu/taxa/species/hygropetra_ balfouriana [Accessed 9 January 2012].
Krammer K. 2000. The genus Pinnularia. In: Diatoms of Europe:diatoms of the European inland waters and comparable habi-tats (Ed. by H. Lange-Bertalot), p. 206. Volume 1. A.R.G.Gantner Verlag K.G., Ruggell. 703 pp.
Lange-Bertalot H. 1997. Frankophila, Mayamaea und Fis-tulifera: drei neue Gattungen der Klasse Bacillariophyceae.Archiv für Protistenkunde 148: 65–76.
Lange-Bertalot H. & Le Cohu R. 1985. Raphe like vestigesin the pennate diatom suborder Araphidineae? Annales deLimnologie 21: 213–220.
Le Cohu R. 1999. Révision des principales espèces de Fragilari-ales (Bacillariophyta) des îles Kerguelen. Canadian Journalof Botany 77: 821–834.
Lowe R.L., Morales E. & Kilroy C. 2006. Frankophilabiggsii (Bacillariophyceae), a new diatom species from NewZealand. Journal of Botany 44: 41–46.
Maidana N.I. & Seeligmann C. 2006. Diatomeas (Bacillario-phyceae) de ambientes acuáticos de altura de la Provincia deCatamarca, Argentina II. Bulletin of the Botanical Society ofArgentina 41: 1–13.
Manguin É. 1964. Contribution à la connaissance des diatoméesdes Andes du Pérou. Mémoires du Muséum Nationald’Histoire Naturelle, Série B, Botanique 12: 41–93.
Mayama S. & Idei M. 2009. Fine structure of two Hygropetraspecies, Hygropetra gelasina sp. nov. and Hygropetra bal-fouriana (Bacillariophyceae), and the taxonomic position ofthe genus with special reference to Frankophila. Phycologi-cal Research 57: 290–298.
McClintic A.S., Casamatta D.A. & Vis M.L. 2003. A sur-vey of algae from montane cloud forest and alpine streamsin Bolivia: macroalgae and associated microalgae. NovaHedwigia 76: 363–379.
Metzeltin D. & Garcıa-Rodrıguez F. 2003. Las DiatomeasUruguayas. Facultad de Ciencias, Universidad de laRepública. Montevideo, Uruguay, Mastergraf S.R.L. 207 p.
Metzeltin D., Lange-Bertalot H. & García-Rodríguez F.
2005. Diatoms of Uruguay. Iconographia Diatomologica 15:1–736.
Morales E.A. 2003. Eighth NAWQA taxonomy workshop on har-monization of algal taxonomy October 25–27. Report No.03-12. Patrick Center for Environmental Research, Academyof Natural Sciences, Philadelphia. 90 pp.
Morales E.A., Vis M., Fernández E. & Kociolek J.P. 2007.Epilithic diatoms (Bacillariophyta) from cloud forest and
alpine streams in Bolivia, South America II: a preliminaryreport on the diatoms from Sorata, Department of La Pax.Acta Nova 3: 680–696.
Morales E.A., Fernández E. & Kociolek P.J. 2009.Epilithic diatoms (Bacillariophyta) from cloud forest andalpine streams in Bolivia, South America 3: diatomsfrom Sehuencas, Carrasco National Park, Department ofCochabamba. Acta Botanica Croatica 68: 263–283.
Patterson B.D., Stotz D.F., Solari S., Fitzpatrick J.W. &Pacheco V. 1998. Contrasting patterns of elevational zona-tion for birds and mammals in the Andes of southeasternPeru. Journal of Biogeography 25: 593–607.
Postek M.T., Howard K.S., Johnson A.H. & McMichael
K.L. 1980. Scanning electron microscopy. A student’shandbook. Ladd Research Industries Inc., Burlington.305 pp.
Ross R., Cox E.J., Karayeva N.I., Mann D.G., Paddock T.B.B.,Simonsen R. & Sims P.A. 1979. An amended terminology forthe siliceous components of thediatom cell. Beihefte zur NovaHedwigia 64: 513–533.
Rott E., Cantonati M., Füreder L. & Pfister P. 2006. Ben-thic algae in high altitude streams of the Alps – a neglectedcomponent of the aquatic biota. Hydrobiologia 562:195–216.
Round F.E. & Bukhtiyarova L. 1996. Four new gen-era based on Achnanthes (Achnanthidium) together witha re-definition of Achnanthidium. Diatom Research 11:345–361.
Round F.E., Crawford R.M. & Mann D.G. 1990. Diatoms. Biol-ogy and morphology of the genera. Cambridge UniversityPress, Cambridge. 747 pp.
Rumrich U., Lange-Bertalot H. & Rumrich M. 2000.Diatomeen der Anden. Von Venezuela bis Patagonien/
Feuerland. Iconographia Diatomologica 9: 1–673.Urrutia R. & Vuille M. 2009. Climate change projections for
the tropical Andes using a regional climate model: tem-perature and precipitation simulations for the end of the21st century. Journal of Geophysical Research 114:D02108,doi:10.1029/2008JD11021.
Van de Vijver B. & Beyens L. 1999. Freshwater diatoms fromIle de la Possession (Crozet Archipelago, sub-Antarctica): anecological assessment. Polar Biology 22: 178–188.
Van de Vijver B., Gremmen N., Beyens L. & Le Cohu R. 2004.Pinnularia sofia Van de Vijver & Le Cohu spec. nov., anew spine-bearing, chain-forming Pinnularia species fromthe sub-Antarctic region. Diatom Research 19: 103–114.
Van de Vijver B., Gremmen N. & Smith V. 2008. Diatomcommunities from the sub-Antarctic Prince Edward Islands:diversity and distribution patterns. Polar Biology 31:795–808.
Van de Vijver B., Agius J.T., Gibson J.A.E. & Quesada A.2009. An unusual spine-bearing Pinnularia species from theAntarctic Livingston Island (South Shetland Islands). DiatomResearch 24: 431–441.
von Stosch H.A. 1975. An amended terminology of the diatomgirdle. Beihefte zur Nova Hedwigia 93: 1–35.
Vuille M., Bradley R.S., Werner M. & Keimig F. 2003. 20thcentury climate change in the tropical Andes: observationsand model results. Climatic Change 59: 75–99.
Young K.R. & León B. 2000. Biodiversity conservation inPeru’s eastern montane forests. Mountain Research andDevelopment 20: 208–211.
