Western North American Naturalist Western North American Naturalist

Volume 73 Number 2 Article 3

7-5-2013

Discovery of the rare freshwater brown alga Discovery of the rare freshwater brown alga Pleurocladia lacustris Pleurocladia lacustris

(Ectocarpales, Phaeophyceae) in California streams (Ectocarpales, Phaeophyceae) in California streams

John D. Wehr Fordham University, Armonk, NY, wehr@fordham.edu

Rosalina Stancheva California State University, San Marcos, CA, rhristov@csusm.edu

Kam Truhn Fordham University, Armonk, NY, truhn@fordham.edu

Robert G. Sheath California State University, San Marcos, CA, rsheath@csusm.edu

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Recommended Citation Recommended Citation Wehr, John D.; Stancheva, Rosalina; Truhn, Kam; and Sheath, Robert G. (2013) "Discovery of the rare freshwater brown alga Pleurocladia lacustris (Ectocarpales, Phaeophyceae) in California streams," Western North American Naturalist: Vol. 73 : No. 2 , Article 3. Available at: https://scholarsarchive.byu.edu/wnan/vol73/iss2/3

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It has been hypothesized that due to theirsmall size, short generation time, and ease ofdispersal, microbial and algal taxa predomi-nantly have cosmopolitan distributions (Fencheland Finlay 2004, Vanormelingen et al. 2008).But important exceptions exist. The well-rec-ognized freshwater green alga Aegagropila lin-naei Kützing is known on at least 3 continents,yet it exists in disjunct locations and is declin-ing in number due to eutrophication and avery limited potential for dispersal (Boedekeret al. 2010). Other species of freshwater algaehave very specific ecological requirements,making their biogeography similarly restricted.The benthic desmid Oocardium stratum Nägeli,a species known since 1849, is rarely observed

and is regarded as globally rare, possibly dueto its close association with CaCO3 deposi -tion (travertine) in limestone springs (Pente-cost 1991, Rott et al. 2010). In recent years,evidence has been growing that other fresh-water algae have limited distributions. Severalare regional endemics with declining popula-tions sufficient to receive conservation status(Kilroy et al. 2007, Cotterill et al. 2008). Thefactors leading to such declines are varied butinclude specialized ecological requirementsand loss of habitat, which can lead to furtherisolated and rare taxa (Sherwood et al. 2004,Coesel and Krienitz 2008). In other cases,reports of algal endemism may be the result ofeither under-sampling or taxonomic confusion

Western North American Naturalist 73(2), © 2013, pp. 148–157

DISCOVERY OF THE RARE FRESHWATER BROWN ALGA PLEUROCLADIALACUSTRIS (ECTOCARPALES, PHAEOPHYCEAE)

IN CALIFORNIA STREAMS

John D. Wehr1,3, Rosalina Stancheva2, Kam Truhn1, and Robert G. Sheath2

ABSTRACT.—Pleurocladia lacustris A. Braun is a freshwater member of the Phaeophyceae, a class of algae that occursalmost entirely in marine waters. It has previously been reported from only about 13 freshwater locations worldwide,just 2 of which are in North America. Outside of North America, P. lacustris has been listed as a threatened species onseveral European red lists. In this paper, we report the discovery of P. lacustris in 3 calcareous streams draining theSanta Lucia Mountains in coastal California, sites that are more than 1200 km from the nearest known population. Pleu-rocladia lacustris is a filamentous, benthic species that forms distinctive hemispherical colonies. It co-occurs with thegreen alga Cladophora glomerata and species of cyanobacteria (Rivularia, Nostoc, Schizothrix spp.). Detailed color illus-trations of the diagnostic macroscopic and microscopic features are provided and appear identical to those features ofEuropean populations. In the California streams, P. lacustris and other associated algae co-precipitate CaCO3 to formcarbonate crusts on rocks. Preliminary ecological data are consistent with other streams and lakes in Europe where P.lacustris has also been reported (pH > 8.0, calcareous substrata, travertine). The global distribution of this presumedrare alga is also described and examined with respect to a specialized ecological niche.

RESUMEN.—Pleurocladia lacustris A. Braun es un organismo de agua dulce perteneciente a Phaeophyceae, una clasede algas que habita casi exclusivamente aguas marinas. Ha sido reportada solo en aproximadamente trece sitios de aguadulce en todo el mundo y sólo dos sitios en los Estados Unidos. Fuera de Norteamérica, está incluida en varias listasrojas de Europa como especie amenazada. En este trabajo presentamos el hallazgo de P. lacustris en 3 arroyos con mate-rial calcáreo que confluyen en las Montañas de Santa Lucía en la costa de California y se encuentran a más de 1200 kmde la población conocida más cercana. Es una especie filamentosa, bentónica, que forma colonias hemisféricas distinti-vas y coexiste con el alga verde Cladophora glomerata y especies de cianobacterias (Rivularia, Nostoc, Schizothrix spp.).Mostramos ilustraciones detalladas en color de las características macroscópicas y microscópicas, donde se puede apre-ciar que son idénticas a las poblaciones europeas. En los arroyos de California, P. lacustris y otras algas asociadas precip-itan CaCO3 y forman cortezas de carbono sobre las rocas. La información ecológica preliminar coincide con la de otrosarroyos y lagos de Europa donde se registró la presencia de esta alga (pH > 8.0, sustrato calcáreo, travertino). Además,se describe y se estudia la distribución global de esta alga aparentemente poco frecuente en relación con un nichoecológico específico.

1Louis Calder Center–Biological Field Station and Department of Biological Sciences, Fordham University, Armonk, NY 10504.2Department of Biological Sciences, California State University, San Marcos, CA 92096.3E-mail: wehr@fordham.edu

148

(Whitford 1983, Vanormelingen et al. 2008).The discovery of new populations of a pre-sumed rare algal species may help to informthis discussion.

The Phaeophyceae is a large and diverseclass of algae with roughly 270 genera and up -wards of 2000 taxa, of which more than 99%occur in marine waters (Wehr 2003). They rangein size from microscopic filaments to largekelps that form huge underwater for ests. Thefreshwater members of this class are modestin size, ranging from small filamentous tufts topseudoparenchymatous crusts (united prostratefilaments), which may form macroscopicallyrecognizable forms in streams and rivers (Kusel-Fetzmann 1996). Few freshwater species arecommonly observed. The crust-forming Herib-audiella fluviatilis (Areschoug) Svedelius is themost widely recognized spe cies, with a fewhundred populations identified from many coun -tries, although thus far only in the northernhemisphere (Wehr 2003). One of the lesser-known freshwater representatives is Pleuro-cladia lacustris A. Braun, a fila mentous speciesdescribed more than 150 years ago (Braun1855), yet observed only from few locations.Interestingly, P. lacustris has been reportedfrom widely separated calcareous freshwaterenvironments (lakes and streams) and also sev-

eral intermittently marine or brack ish loca-tions (Eloranta et al. 2011). Some freshwaterpopulations have apparently been extirpateddue to habitat degradation and urbanization(Sukopp 2003, Täuscher 2011, Wehr 2011),causing some European countries to list thisspecies on their conservation red lists (Sie -mińska 2006, Täuscher 2010). In this paper,we report the recent discovery of P. lacustrisin 3 streams in California. These streams con-stitute the fourth location for North America(third freshwater location), more than 1200 kmfrom the nearest known population. We alsoexamine the distribution of this species globallyand discuss the presumed rarity of its occur-rence (e.g., Siemińska 2006, Täuscher 2011).

STUDY AREA

The study was conducted along the Big Surcoast in central California (Fig. 1). A previouscollection of periphyton taken from a streamin this area on 23 June 2010 by the SurfaceWater Ambient Monitoring Program (SWAMP)of the California State Water Resources ControlBoard included material that was thought tobe P. lacustris. In the present study, we sur-veyed 12 streams that drain the Santa LuciaMountains between Pfeiffer Big Sur State Parkand San Simeon, California. Most are first- orsecond-order streams and typically have verysteep gradients with a calcareous geology (Hen -son and Usner 1993); CaCO3 precipitationand travertine formation are apparent at manylocations. The region has a cool-Mediterraneanclimate, and riparian vegetation is composedmainly of white alder (Alnus rhom bifolia), big -leaf maple (Acer macrophyllum), arroyo willow(Salix laseolepis), black cottonwood (Populustrichocarpa), California bay (Umbellularia cali-fornica), and in some locations, coast redwood(Sequoia sempervirens). The streambed at mostsites had variable canopy cover, with areasranging from open sun to heavy shade. Fur-ther details of the physical conditions of thesestreams are described by Rundio (2009).

METHODS

Stream sites were accessed from along a40-km stretch of California State Route 1, northfrom Cambria, California, 12–14 March 2012.All sampling reaches were located at least 150m upstream of any marine water. We collectedalgae by using knives and razor blades to

2013] FRESHWATER PLEUROCLADIA IN CALIFORNIA 149

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Fig. 1. Location of 3 stream populations of Pleurocladialacustris (solid circles), with map of study area in westcentral California (inset).

sample rocks, and we inspected the materialwith a Swift FM-31 field microscope. At siteswhere P. lacustris was present, samples werecollected into bags or tubes (at least 10 repli-cates per site) and separated for future use inmi crophotography, lab cultures, DNA extrac-tion, and preserved samples. Live materialwas placed in sterile, 120-mL Whirl-Pak® bagsand stored on ice. Water temperature and spe-cific conductance were measured in situ byusing a YSI EC300 meter, and pH was mea-sured by using a YSI pH10A meter. A portablelaboratory microscope (Nikon 50i with LEDillumination) was later used after each sam-pling day to confirm identifications conductedin the field.

Samples were shipped overnight to ourlaboratories for further observations and mi -crophotography. In the laboratory, live algalsamples were prepared for wet-mount micro -scopy by using sterile probes to separate colo -nies prior to observation. Occasionally, diluteacid (1% HCl) was added to heavily calcifiedcolonies to improve clarity. Specimens wereobserved using a Nikon Eclipse E600 micro-scope with Nomarski (DIC) optics (200X, 400X,and 1000X total magnification), photographedusing a Nikon DS-Fi1 digital camera, andprocessed and measured using NIS-Elementssoftware. Identifications were based on Kusel-Fetzmann (1996), Wehr (2003), and Elorantaet al. (2011). Georeferenced voucher specimenswere submitted to the William and LyndaSteere Herbarium at the New York BotanicalGarden (NYBG), and DNA samples werestored frozen (–40 °C) at the NYBG MolecularSystematics Laboratory for a later study. Smallportions of live material were placed in algalgrowth media and maintained at FordhamUniversity and the NYBG.

Past records of Pleurocladia lacustris werecollected from published literature, personalcommunications with authors, and herbariumrecords. Where precise georeferenced records

were not available, latitude and longitude val-ues for the names or nearest water body wereused. Geographic data for all populations werecompiled and mapped using ArcGIS v. 10.

RESULTS

Of 7 streams sampled, we located 3 thatcontained populations of Pleurocladia lacustris(Table 1). Water at all sites was alkaline (pH>8.0) and had moderate specific conductance(average ~250 mS ⋅ cm–1). When present, thisalga was among the most common and macro-scopically recognizable species in the stream.Along with Pleurocladia, the filamentous greenalga Cladophora glomerata and colonial cyano -bacteria Nostoc, Rivularia, and Schizothrixspecies were also common. Macroscopically,the morphology of Pleurocladia appeared tohave 2 distinct forms: small irregular, flatbrown spots on rocks (Fig 2A) and brown totan-colored hemispherical tufts (Fig. 2B). Bothforms occurred in running water, but thehemispherical form was more common in low-current microhabitats, whereas the spreading(flat-form) was more common in riffles. In thefield, colonies of both forms ranged in sizefrom about 1 to 5 mm in diameter and ap -peared dark brown or reddish brown in color.Where colonies were heavily calcified, theyappeared light tan in color. Those that pro-duced large quantities of elongate hyaline hairs(narrow, unpigmented filaments), grayish mats,or tufts were also apparent.

Microscopically, basal filaments were alluniaxial and ranged in diameter from 12 to 20mm, whereas upright filaments were 9–14 mmwide (Figs. 2C–D). Branching frequency wasirregular, although filaments in larger cush -ion-like colonies tended to be more elongateand have fewer side branches. In smaller oryounger colonies, the typical comb-like ap -pearance was common. In both forms, cells inthe main axes ranged from 10 to 35 mm in

150 WESTERN NORTH AMERICAN NATURALIST [Volume 73

TABLE 1. Locations and ecological conditions of California streams from which Pleurocladia lacustris was collected in2012.

Stream Salmon Creek Willow Creek Villa Creek

Latitude (N) 35°48�56.80� 35°53�37.23� 35°50�59.24�Longitude (W) 121°21�31.82� 121°27�40.71� 121°24�28.23�Date 12 Mar 2012 12 Mar 2012 13 Mar 2012Temperature (°C) 9.3 11.4 10.9pH 8.22 8.45 8.50Specific conductance (mS ⋅ cm−1) 220.9 253.2 271.2

2013] FRESHWATER PLEUROCLADIA IN CALIFORNIA 151

Fig. 2. Morphology of field populations of Pleurocladia lacustris from California streams: A, macroscopic appearanceof dark brown, spreading (flat-form) colonies on rocks; B, macroscopic appearance of cushion-forming colonies (grayishmaterial is a network of elongate hyaline cells); C, microscopic appearance of cushion-forming colonies with terminalhyaline cells (arrows); D, microscopic appearance of spreading colonies, with unilocular sporangia (arrows) and CaCO3encrusting the colony base (brackets); E, detail of filaments with single parietal chloroplast and numerous oil bodies;F, detail of unilocular sporangium (left) on lateral side branch.

length. Chloroplasts were single and parietal,usually filling one-half to three-quarters of thecell diameter (Fig. 2E). In older cells, somechloroplasts appeared twisted or lobed. Nu -merous spherical bodies, assumed to be oildroplets, were scattered throughout the cells.When present, terminal hyaline hair cells (Fig.2C) often would exceed 200 mm in length.Macroscopically, some appeared to extend morethan 1 mm. Large, ovoid, or clavate unilocularsporangia (20–40 × 30–85 mm) were frequentlyformed on short side branches (Fig. 2F). Pluri -locular sporangia were not observed.

Following the discoveries in California, theglobal distribution of Pleurocladia lacustrisnow includes 8 countries, and 4 locations inNorth America, totaling 27 separate freshwa-ter stream or lake sites. Most sites are concen-trated in western and northern Europe (Fig. 3,Appendix), and many of these are located ineastern Sweden, including several rivers inthe eastern Uppland province north of Stock-holm and various standing waters in the Öre-grund Archipelago area (Israelsson 1938, Waern1952). Ecological information from these re -ports is spotty, but many researchers commenton the calcareous nature of P. lacustris. Of allthe freshwater populations currently known,the California sites are nearest to a populationin the Green River, Utah, downstream ofFlaming Gorge Dam (Ekenstam et al. 1996,Wehr 2003). These 2 locations are more than

1200 km apart, but the macroscopic morphol-ogy of the 2 populations is indistinguishable interms of cell dimensions, branching pattern,and sporangia. Both occur in calcareous river-ine habitats and commonly have CaCO3 en -crusting at the base of the colonies (Fig. 2D).One recent collection has also been madefrom a deep limestone pool in South Australia(deposited in the National Herbarium of NewSouth Wales; listed as Pleurocladia sp. nov.but without verification; #NSW 486949), asite which is ~13,000 km from the Californiapopulations and ~15,000 km from the nearestfreshwater population in Europe.

DISCUSSION

The evidence for the existence of rare spe -cies of freshwater algae and other protists hasbeen regarded somewhat skeptically by someresearchers (Whitford 1983, Fenchel and Fin-lay 2004). Reports of restricted distributions infreshwater algae may simply be the result ofundersampling (Foissner 2008). Some speciesare seasonally ephemeral and missed by spotsampling. Others have cryptic life stages thatroutine surveys may miss. However, studiesof freshwater algae in North America havespanned more than 150 years, and certain spe -cies are still rarely seen. Evidence from multi-ple studies in Europe suggest that there arerare, threatened, and endangered species of

152 WESTERN NORTH AMERICAN NATURALIST [Volume 73

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Fig. 3. Geographic locations of all known populations of Pleurocladia lacustris (solid circles). Details of individualsites, including designations of freshwater, brackish, and marine locations, are given in the Appendix.

algae that require conservation efforts (Brodieet al. 2007, Cotterill et al. 2008, Ros et al.2009). This idea has been raised specificallyfor Pleurocladia lacustris in the United King-dom, where the only known habitats havebeen altered through nutrient pollution (Brodieet al. 2007) and the species may have beenregionally extirpated (Wehr 2011). Similarly,Friedrich et al. (1984) documented that all 4species of freshwater brown algae known fromGermany are listed as endangered to differentdegrees. This includes the population of Pleu-rocladia lacustris from its type location in Ger-many, where it was first described by Braun in1855 and where it is now listed as threatenedwith local extinction (Geissler 1988). As of thisdate, it is not known if this important popula-tion still exists.

The ecological data for P. lacustris assem-bled so far indicate that the species is re -stricted to fairly nutrient-rich, alkaline, cal-careous environments (Israelsson 1938, Waern1952, Kusel-Fetzmann 1996) and that it isabsent from softwater or humic-stained sys-tems (Kann 1940, Eloranta et al. 2011). A sin-gle sample in June 2010 taken from one of oursites (Salmon Creek) by the Surface WaterAmbient Monitoring Program of the CaliforniaState Water Resources Control Board indi-cated moderately high specific conductance(372 mS ⋅ cm–1), very low TDP (9.5 mg ⋅ L–1),TDN 63.6 mg ⋅ L–1, and alkaline pH (8.5) butclearly nonsaline. These data collectively indi-cate that the alga occurs in systems with pH>7.5 and specific conductivity >200 mS ⋅cm–1. No extensive analyses of water chem-istry requirements exist yet for this species.However, preliminary lab experiments suggestthat P. lacustris may tolerate a wide range ofphosphorus conditions through the productionof hyaline hairs (Wehr 2003). Hyaline hairs inother species have been documented to utilizeorganic-P sources (Gibson and Whitton 1987).Pleurocladia lacus tris has been collected fromsmall ponds to very large lakes (e.g., Klebahn1895, Kirkby et al. 1972, Kahlert et al. 2002,Young et al. 2010), as well as from streams andrivers (Eken stam et al. 1996, Kusel-Fetzmann1996, this study).

Interestingly, in addition to occurring in anumber of fully freshwater systems, P. lacus-tris has been reported from brackish environ-ments, often in areas where limestone is pres -ent (Waern 1952, Wilce 1966, Konan-Brou and

Guiral 1994, Aysel et al. 2008). The specificconductance data from the 3 California sites(220–270 mS ⋅ cm–1) indicate that these popu-lations have little or no marine influence. Thespecies is strictly benthic in habit but growson a variety of substrata, with several reportsof occurrence on stones (Kann 1993, Kusel-Fetzmann 1996, Kahlert et al. 2002), as an epi-phyte on higher plants or other algae (Kirkbyet al. 1972, Szymanska and Zakrys 1990, Younget al. 2010), and even as an endophyte in a fewinstances (Waern 1952). Kann and Tschamler(1976) demonstrated that the alga also colo-nizes artificial substrata (plastic and glassslides) in suitable habitats. Given these eco-logical limitations, this species may be geo-graphically limited by a restriction to calcare-ous habitats.

Pleurocladia lacustris was the first freshwa-ter species of brown algae described; yet it isan enigma with regard to its ecological re -quirements and geographic distribution. Withfreshwater and brackish water populations,and relatively few known locations, the dis-covery of new populations is noteworthy.Further biodiversity surveys may clarify theextent to which P. lacustris may be a rarespecies. But, given its widespread locationsseparated by long distances, 4 broad questionsemerge: (1) Is “Pleurocladia lacustris,” as re -ported, one species or several? It may be thatwhile its morphological characteristics arehighly consistent among populations, isola -tion and divergence may have resulted in sev-eral cryptic species, as demonstrated in otherfreshwater and marine taxa (Casamatta et al.2003, Saunders 2008). Future genetic analyseswill shed light on this question. (2) If its distri-bution is restricted, is P. lacustris limited by itsecological requirements or perhaps by disper-sal? Field data suggest that this species pre -fers calcareous conditions, but detailed experi-ments are needed to confirm either factor as acause of its ecological and geographic restric-tion (e.g., Telford et al. 2006, Boedeker et al.2010). (3) Is P. lacustris a rare species or eco-logically restricted? Though habitat destruc-tion has reduced its distribution in some areas,P. lacustris could still be undersampled ormissed in biodiversity surveys. New surveyscan employ molecular methods, such as ge -nome shotgun sequencing and DNA bar-coding, to reveal hidden biological diversityand the presence of underreported taxa (e.g.,

2013] FRESHWATER PLEUROCLADIA IN CALIFORNIA 153

Eisen 2007). (4) What is the relationship be -tween freshwater and brackish (intermittentlymarine) populations of P. lacustris, both interms of ecological adaptations and evolution?Laboratory studies can elucidate the ability ofnear-coastal and inland populations to adapt toa range of salinities (e.g., Dittami et al. 2012),whereas population genetics studies may ex -plain the evolutionary route this species mayhave taken from marine to freshwater habitats(e.g., Müller et al. 1998).

ACKNOWLEDGMENTS

We thank the Surface Water Ambient Moni -toring Program (SWAMP)–California State Wa -ter Resources Control Board for providing theinitial periphyton sample containing Pleuro-cladia. We also thank Christina Vanderwerkenfor her enthusiastic and capable assistancewith fieldwork, as well as Professor RobertWilce (University of Massachusetts) for adviceon the biology and distribution of Pleurocla-dia. JDW was supported in part by a FacultyResearch Grant from Fordham University. RGSand RS were supported by the California StateWater Resources Control Board–SWAMP.

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156 WESTERN NORTH AMERICAN NATURALIST [Volume 73A

PPE

ND

IX. L

istin

g of

all

know

n po

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tions

of P

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2013] FRESHWATER PLEUROCLADIA IN CALIFORNIA 157A

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