- Email: [email protected]
New records of Vaucheria species (Xanthophyceae) with associated Proales werneckii (Rotifera) from North America
cvptogamie,
67
Algol., 1999, 20 (2): 67-73
New records of kucheriu species (Xanthophyceae) with associated Proales werneckii (Rotifera) from North America Robert
G. VERB”,
Morgan L. VI,!?, Donald W OT? & Robert L. WALLACE’
U Environmental & Plant Biology Department, Ohio University, Athens, Ohio, U.S.A. 45701; [email protected]; [email protected] ’ Department of Biology, The University of Akron, Akron, Ohio, U.S.A. 44325-3908; [email protected] ’ Department of Biology, Ripon College, Ripon, Wisconsin, U.S.A. 54971-0248; [email protected] (Received 17 September 1998, accepted 25 January I999)
Abstract - The presence of galls on species of Vaucheria was investigated both seasonally and in a number of locations in North America. These galls are induced by the rotifer, Proales wemeckii. In an Ohio stream, Vaucheria bursata and V geminata were found to have galls throughout their growing season; September to January. Galls were most abundant in October and ranged in size from 90-260 urn in width and 140-790 urn in length. New records of Vuucheriu with galls from locations in Alabama, California, Georgia, Louisiana, North Carolina, Ohio and Tennessee are reported. Three of the five taxa collected were not previously known to contain galls. In addition, worldwide literature on the distribution of these associated taxa was reviewed. 0 ADAC / Elsevier, Paris freshwater algae / new records I North America Rotifera I Vaucheria / Xanthophyceae
1 Ohio I parasitism
/ Proales werneckii
I
RCsumC - La presence de vesicules sur des especes de Vaucheria a BtC examinee durant l’annde et dans plusieurs localites d’Am&ique du Nord. Ces vesicules sont provoquees par le rotifere parasite, Proales werneckii. Dans une rivitre de I’Ohio, Vaucheria bursata et V: geminata ont CtC trou& avec des vesicules durant les mois de septembre a janvier, les mois durant lesquels ces especes de Gzaucheria se developpent dans cette riviere. Les vesicules sont plus nombreuses en octobre. Leurs dimensions vont de 90 a 260 urn en largeur et de 140 a 790 urn en longueur. De nouveaux cas de Vaucheria avec ces vesicules ont t%e signal& en Alabama, Califomie, Caroline du Nord, GCorgie, Louisiane, Ohio et Tennessee. Chez trois des cinq especes examinees, la ptisence de galles dues a la presence du rotifere n’avait pas encore ettCsignaXe. De plus, une synthbse de la literature relative a la distribution de ces espbces est prcsentee. 0 ADAC I Elsevier, Paris algues d’eau deuce I Amkrique Vaucheria / Xanthophyceae
du Nord / Ohio / parasitisme
I Proales werneckii
I rotiEre
INTRODUCTION
worldwide.
The xanthophyte genus, Vaucheria, occurs in marine, estuarine and freshwaters numerous habitat types, including moist Freshwater species colonize
* Correspondence
and reprints
I
R.G. Verb, M.L. Vis, D.W. Ott & R.L. Wallace
68
Fig. 1. A mature Kz’aucheria gall containing Proales werneckii (double arrowheads) and eggs (single arrowhead) from un-named tributary in the Hocking River drainagebasin,southeastern Ohio. , Fig. 2. A Vuucheriu geminata gametophore induced in culture from unnamed tributary in the Hocking River drainagebasin,southeastern Ohio. Fig. 3. Reproductive structures of tiaucheria bursata induced in culture from unnamed tributary in the Hocking River drainage basin, southeastern Ohio.
soils, shallow ditches and the edges of lakes and streams(Entwisle, 1988). Typically Vaucheria speciesform a densemacroscopic mat of intertwined coenocytic filaments which may or may not be reproductive at time of collection. Vaucheria specieshave been reported to be parasitized by the rotifer Proales werneckii Ehrenberg (Tab. 1). Once inside the alga, the 140-200 pm rotifer induces the formation of a conspicuousgall and consumescytoplasm and cytoplasmic organellesof the alga (Ott, 1977) (Fig. I). The female may deposit upwards of 80 eggs inside the gall. The eggs may hatch in the sameseasonand the young rotifers escapefrom the gall to searchfor new filaments to parasitize or the eggs overwinter in the sedimentsand hatch in the following season(De Smet, 1996). Although the Kzucheria sp. and Proales werneckii associationis well known, there have been only scattered notes in the literature on the occurrence of these galls (e.g. Christensen, 1969, 1987; Entwisle, 1988). Davis & Gworek (1973) reported the associationbetween Vaucheria sp. and Proales werneckii in a collection from a Florida spring that was the first well-documented site in North America. Habeeb (1965) mistakenly described these galls as reproductive structures and proposed a new taxon, Debsalga gigasporungia, for a collection from California. In addition, there is little information on the seasonaloccurrence of this association. Therefore, this study was undertaken to provide additional information on the seasonalityof this relationship and new localities in North America.
MATERIALS
AND METHODS
Seasonal Study An un-namedsecondorder tributary of the Hocking River in southeasternOhio, USA
(N 39O24.197’,
W 82O12.993’)
was visited
on a monthly
basis from
May
1997
Records
Tab. 1. Reports
with
location
of Vaucheria
with Proales
and associated
Proales
uversa Hassall (0. F. Miiller) C. A.
(L.) Christensen
V dichofoma (L.) C. A. Agardh V dillwynii (Weber & Mohr) C. A. Agardh
Netherlands Firestone Metropolitan Park, Akron, OH, U.S.A. Athens County, OH, U.S.A. Switzerland Australia
Switzerland Australia Netherlands Netherlands Australia Netherlands
V discoidea Taft I! erythrospora Christensen V frigida (Roth) C. A. Agardh V geminata (Vaucher) De Candolle
k! racemosa
V prona
De Candolle
Christensen
V terresfris Vaucheria
(Vaucher)
(Vaucher)
werneckii.
Reference
Netherlands V canalicularis
69
Location’
Species Vaucheria V bursata Agardh
of Vaucheria
De Candolle
sp.
’ AL = Alabama, CA = California, OH = Ohio, TN = Tennessee.
Green, OH, U.S.A. Australia Switzerland Athens County, OH, U.S.A. Firestone Metropolitan Park, Akron, OH, U.S.A. Ponchatoula, LA, U.S.A. Woodstock, GA, U.S.A. Santa Cruz, CA, U.S.A. Switzerland Australia Netherlands
Firestone Metropolitan Park, Akron, OH, U.S.A. Nimisila Reservoir, Akron, OH, U.S.A. Hammond, LA, U.S.A. Ponchatoula, LA, U.S.A. Nashville, TN, U.S.A. Chapel Hill, NC, U.S.A. Richfield, OH, U.S.A. Carrboro, NC, U.S.A. Lauderdale County, AL, U.S.A. United Kingdom Switzerland Florida. U.S.A. GA = Georgia,
Simons & Vroman This study This study Christensen (1969) Cribb (1964, as 1! Entwisle (1988) Simons & Vroman v sessilis) Christensen (1969) Christensen (1987) Simons & Vroman I! woroniniana) Simons & Vroman Christensen (1987)
(1968)
se&is), (1968,
as
(1968, as (1968)
Simons & Vroman (1968, as V pachydema) This study Christensen (1987) Christensen (1969) This study This study This study This study This study Christensen (1969) Christensen (1987) Simons & Vroman (1968, as t! walzi), Prud’homme van Reine (1970) This study This study This study This study This study This study This study This study This study Rousselet (1897) Christensen (1969) Davis & Gworek (1973)
LA = Louisiana,
NC = North
Carolina,
70
R.G. Verb, M.L. Vis, D.W. Ott & R.L. Wallace
through May 1998. Samples of macroscopic algae were collected and fixed in 2.5 % calcium carbonate-buffered glutaraldehyde. Maximum width, average depth, pH, specific conductance, temperature and mean current velocity were measured as described by Sheath et al. (1989). In addition, water samples were analyzed for orthophosphate, nitrate, total iron, aluminum and manganese with a DR/890 calorimeter, using standard protocols and powder pills from Hach Company (Anonymous, 1997). Water turbidity was measured with a Hach 21OOP’” turbidity meter. Species identifications of Kzucheria were determined by inducing gametophore production. Fresh material from the field was cultured in stream water for one week in a diurnal growth chamber at 15-20” C with fluorescent lighting providing 2OwOO lux under a 12-12 h photoregime. Species identification and nomenclature follows Rieth (1980) and Entwisle (1988). The rotifer was identified and nomenclature follows De Smet (1996). The length and width of galls containing~animals were measured using an 0~~1% micrometer. Images were taken with a Pixera digital camera attached to a BX40 Olympus microscope. A voucher specimen of the Kzucheria with galls has been deposited at the Ohio State University Herbarium (OS) under the accession number OS-399245. Collections
from North America
Kzucheria samples were collected in a number of locations from eastern North America and California (Tab. 1). These collections have been made sporadically over the past 30 years and some from the same location repeatedly. Sterile thalli with galls were transported back to the laboratory and placed in finger bowls with water from the collection site. The finger bowls were placed in a growth chamber at 12-15” C with fluorescent lighting providing 2000-4000 lux under a 14-10 hour photoregime. Cultures were left in the growth chamber for two weeks, after which time gametophore development had occurred and species could be identified. Unfortunately, no material was available for deposit in the Ohio State University Herbarium.
RESULTS
AND DISCUSSION
Seasonal Study Prior to September 1997 (May-August 1997), no visible Vuucheriu thalli had been collected from the Hocking River tributary location. In September 1997, Vaucheriu was found growing as a conspicuous mat in a large stream pool. Upon microscopic examination of Kzucheriu thalli, galls were observed (Fig. 1). The organism living in the galls was identified as the rotifer Proules werneckii. Vaucheriu with Proales wemeckii was present in stream collections from September 1997 through January 1998. Proales galls were most abundant in the October 1997 sample, but galls were present during all five months. Neither Kzucheriu nor associated Proales werneckii were collected in February 1998 or subsequent collections (March-May 1998). Therefore, it appears Proales wemeckii rapidly infects Vaucheriu mats since the first appearance of the thalli already shows gall formation. In addition, Proales is present throughout the tiucheriu growing season. Throughout the five sampling months, the Vaucheriu specimens collected were vegetative, lacking gametophores. Vaucheria specimens from the September sampling
Records of Vaucheria with Proales
71
were induced to form gametophores in culture and identified as two distinct species, Vaucheria geminata and K bursata (Figs 2, 3). Both species were colonized by Proales werneckii, possibly demonstrating lack of species specific infection. Measurements of width and length for 56 galls containing Proales wemeckii from the October sampling varied considerably. Galls ranged from 90-260 pm in width and 140-790 pm in length with a mean gall size of 135 x 320 pm. This wide range in gall dimensions may be due to progressive stages of gall development. Stream conditions were monitored throughout the sampling period (May 1997May 1998). The associated Kzucheria and Proales werneckii were able to tolerate a wide range in stream temperature, which fluctuated from 4 to 23” C. However, they are only found in association for a short period of time in the fall to early winter. During this time stream pH was neutral. Specific conductance was high (680-794 @cm-‘) for the region probably due to dissolved metals (Fe, Al and Mn). Orthophosphate levels were high and probably caused by a variety of anthropogenic inputs such as leachates from septic tanks and refuse present along the edge of the stream. These results suggest that Kzucheria bursata, V. geminuta and Proales wemeckii have a high tolerance of both organic and inorganic pollution. The period during which tiucheria with Proales was collected did not appear to have stream conditions uncharacteristic for the remainder of the sampling time.
Collections from North America Kzucheria species from California on the Pacific coast and Alabama, Georgia, Louisiana, North Carolina, Ohio and Tennessee in the eastern US were collected with Proales wemeckii galls (Tab. 1). These collections represent 13 new localities from states which, for the most part, had no previous records of this association. In North America, these associated taxa have only been reported from one Florida spring location (Davis & Gworek, 1973), California (as Debsalga gigasporangiu) (Habeeb, 1965) and noted to occur with no specific information on location (Wolle, 1887). Our collections confirm that this association is more prevalent in North America than previously thought. These taxa occurring together are probably cosmopolitan, but detailed information on geographical distribution suffers from the lack of widescale investigation. Worldwide, reports of this association have been sporadic with specific sites from Australia (Cribb, 1964; Christensen, 1987; Entwisle, 1988), Switzerland and France (Christensen, 1969), the United Kingdom (Rousselet, 1897) and the Netherlands (Prud’homme van Reine, 1972; Simons & Vroman, 1968) (Tab. 1). Again, when researchers have conducted extensive surveys of particular regions, Pro&s wemeckii in conjunction with Vaucheria species has been found in a number of localities (Christensen, 1969, 1987; Entwisle, 1988). Other authors have made reference to the association, in either literature reviews or early research, hut little information on location was included (Balbiani, 1878; Bock, 1954; Debray, 1890; Dormer, 1966; Ehrenberg, 1834; Hofker, 19 18; Gabriel, 1922; May, 1989; Rieth, 1980; Rothert, 1896; Weidner, 1952; Wolle, 1887). Although information is scarce, it would appear from the literature that this association has been known for quite some time. In this study, five distinct species were identified with Pro&es werneckii galls (Tab. 1). Vaucheria discoidea, V geminata and V pronu are new records of species forming galls and only K bursutu and K terrestris had been previously noted. There are at least 12 Vaucheriu species known worldwide to have associatedProules wemeckii including those reported in this study (Tab. 1). All Vaucheriu speciesreported forming galls are from sectionsComiculutue. Racemosue,T’buligerae, Kzucheria and Woroninia,
Records of Vaucheria with Proales
73
MAY L., 1989 - Epizoic and parasitic rotifers. Hydrobiologia 186/187: 59967. OlT D.W., 1977 - Ultrastructural observations on parasitism of Vaucheria prona by Proales wemeckii. Journal of Phycology 13 (supplement): 5 1. PRUD’HOMME VAN REINE W.F., 1972 - Gallen in Vaucheria-draden (Algen-Xanthophyceae). Gorteria 6: 89-95. RIETH A., 1980 - Xanthophyceae. Part 2. In: Ettl, H., Gerloff, J. and Heynig, H. (eds), Siisswasserfiorft van Mitteleuropa. Vol. 4. Stuttgart, Gustav Fisher Verlag, 147 p. ROTHERT W., 1896 - Uber die Gallen der Rotatorie Notommata wemeckii auf Kzucheria walzi n. sp. Jahrbuecher fuer Wissenschajiliche Botanik 29: 525-594. ROUSSELET CF., 1897 - On the male of Proales wemeckii. Journal. Quekett Microscopical Club, series 2, 6: 415-418. SHEATH R.G., HAMILTON PB., HAMBROOK J.A. & COLE K.M., 1989 - Stream macroalgae of the eastern boreal forest region of North America. Canadian Journal of Botany 58: 1295-1318. SIMONS J. & VROMAN M., 1968. Some remarks on the genus Vaucheria in the Netherlands. Acta Botanica Neerlandica 17: 461-466. WEIDNER H., 1952. Proales wemecki, ein in Vaucheria parasitierendes Rotator. Nuchrichten. Natunvissenschaftliches Museum der Stadt {Aschafenburg) 35: 3946. WOLLE F., 1887 - Fresh-water Algae of the United States. Bethlehem, Pennsylvania, The Comenius Press, 364 p.
67
Algol., 1999, 20 (2): 67-73
New records of kucheriu species (Xanthophyceae) with associated Proales werneckii (Rotifera) from North America Robert
G. VERB”,
Morgan L. VI,!?, Donald W OT? & Robert L. WALLACE’
U Environmental & Plant Biology Department, Ohio University, Athens, Ohio, U.S.A. 45701; [email protected]; [email protected] ’ Department of Biology, The University of Akron, Akron, Ohio, U.S.A. 44325-3908; [email protected] ’ Department of Biology, Ripon College, Ripon, Wisconsin, U.S.A. 54971-0248; [email protected] (Received 17 September 1998, accepted 25 January I999)
Abstract - The presence of galls on species of Vaucheria was investigated both seasonally and in a number of locations in North America. These galls are induced by the rotifer, Proales wemeckii. In an Ohio stream, Vaucheria bursata and V geminata were found to have galls throughout their growing season; September to January. Galls were most abundant in October and ranged in size from 90-260 urn in width and 140-790 urn in length. New records of Vuucheriu with galls from locations in Alabama, California, Georgia, Louisiana, North Carolina, Ohio and Tennessee are reported. Three of the five taxa collected were not previously known to contain galls. In addition, worldwide literature on the distribution of these associated taxa was reviewed. 0 ADAC / Elsevier, Paris freshwater algae / new records I North America Rotifera I Vaucheria / Xanthophyceae
1 Ohio I parasitism
/ Proales werneckii
I
RCsumC - La presence de vesicules sur des especes de Vaucheria a BtC examinee durant l’annde et dans plusieurs localites d’Am&ique du Nord. Ces vesicules sont provoquees par le rotifere parasite, Proales werneckii. Dans une rivitre de I’Ohio, Vaucheria bursata et V: geminata ont CtC trou& avec des vesicules durant les mois de septembre a janvier, les mois durant lesquels ces especes de Gzaucheria se developpent dans cette riviere. Les vesicules sont plus nombreuses en octobre. Leurs dimensions vont de 90 a 260 urn en largeur et de 140 a 790 urn en longueur. De nouveaux cas de Vaucheria avec ces vesicules ont t%e signal& en Alabama, Califomie, Caroline du Nord, GCorgie, Louisiane, Ohio et Tennessee. Chez trois des cinq especes examinees, la ptisence de galles dues a la presence du rotifere n’avait pas encore ettCsignaXe. De plus, une synthbse de la literature relative a la distribution de ces espbces est prcsentee. 0 ADAC I Elsevier, Paris algues d’eau deuce I Amkrique Vaucheria / Xanthophyceae
du Nord / Ohio / parasitisme
I Proales werneckii
I rotiEre
INTRODUCTION
worldwide.
The xanthophyte genus, Vaucheria, occurs in marine, estuarine and freshwaters numerous habitat types, including moist Freshwater species colonize
* Correspondence
and reprints
I
R.G. Verb, M.L. Vis, D.W. Ott & R.L. Wallace
68
Fig. 1. A mature Kz’aucheria gall containing Proales werneckii (double arrowheads) and eggs (single arrowhead) from un-named tributary in the Hocking River drainagebasin,southeastern Ohio. , Fig. 2. A Vuucheriu geminata gametophore induced in culture from unnamed tributary in the Hocking River drainagebasin,southeastern Ohio. Fig. 3. Reproductive structures of tiaucheria bursata induced in culture from unnamed tributary in the Hocking River drainage basin, southeastern Ohio.
soils, shallow ditches and the edges of lakes and streams(Entwisle, 1988). Typically Vaucheria speciesform a densemacroscopic mat of intertwined coenocytic filaments which may or may not be reproductive at time of collection. Vaucheria specieshave been reported to be parasitized by the rotifer Proales werneckii Ehrenberg (Tab. 1). Once inside the alga, the 140-200 pm rotifer induces the formation of a conspicuousgall and consumescytoplasm and cytoplasmic organellesof the alga (Ott, 1977) (Fig. I). The female may deposit upwards of 80 eggs inside the gall. The eggs may hatch in the sameseasonand the young rotifers escapefrom the gall to searchfor new filaments to parasitize or the eggs overwinter in the sedimentsand hatch in the following season(De Smet, 1996). Although the Kzucheria sp. and Proales werneckii associationis well known, there have been only scattered notes in the literature on the occurrence of these galls (e.g. Christensen, 1969, 1987; Entwisle, 1988). Davis & Gworek (1973) reported the associationbetween Vaucheria sp. and Proales werneckii in a collection from a Florida spring that was the first well-documented site in North America. Habeeb (1965) mistakenly described these galls as reproductive structures and proposed a new taxon, Debsalga gigasporungia, for a collection from California. In addition, there is little information on the seasonaloccurrence of this association. Therefore, this study was undertaken to provide additional information on the seasonalityof this relationship and new localities in North America.
MATERIALS
AND METHODS
Seasonal Study An un-namedsecondorder tributary of the Hocking River in southeasternOhio, USA
(N 39O24.197’,
W 82O12.993’)
was visited
on a monthly
basis from
May
1997
Records
Tab. 1. Reports
with
location
of Vaucheria
with Proales
and associated
Proales
uversa Hassall (0. F. Miiller) C. A.
(L.) Christensen
V dichofoma (L.) C. A. Agardh V dillwynii (Weber & Mohr) C. A. Agardh
Netherlands Firestone Metropolitan Park, Akron, OH, U.S.A. Athens County, OH, U.S.A. Switzerland Australia
Switzerland Australia Netherlands Netherlands Australia Netherlands
V discoidea Taft I! erythrospora Christensen V frigida (Roth) C. A. Agardh V geminata (Vaucher) De Candolle
k! racemosa
V prona
De Candolle
Christensen
V terresfris Vaucheria
(Vaucher)
(Vaucher)
werneckii.
Reference
Netherlands V canalicularis
69
Location’
Species Vaucheria V bursata Agardh
of Vaucheria
De Candolle
sp.
’ AL = Alabama, CA = California, OH = Ohio, TN = Tennessee.
Green, OH, U.S.A. Australia Switzerland Athens County, OH, U.S.A. Firestone Metropolitan Park, Akron, OH, U.S.A. Ponchatoula, LA, U.S.A. Woodstock, GA, U.S.A. Santa Cruz, CA, U.S.A. Switzerland Australia Netherlands
Firestone Metropolitan Park, Akron, OH, U.S.A. Nimisila Reservoir, Akron, OH, U.S.A. Hammond, LA, U.S.A. Ponchatoula, LA, U.S.A. Nashville, TN, U.S.A. Chapel Hill, NC, U.S.A. Richfield, OH, U.S.A. Carrboro, NC, U.S.A. Lauderdale County, AL, U.S.A. United Kingdom Switzerland Florida. U.S.A. GA = Georgia,
Simons & Vroman This study This study Christensen (1969) Cribb (1964, as 1! Entwisle (1988) Simons & Vroman v sessilis) Christensen (1969) Christensen (1987) Simons & Vroman I! woroniniana) Simons & Vroman Christensen (1987)
(1968)
se&is), (1968,
as
(1968, as (1968)
Simons & Vroman (1968, as V pachydema) This study Christensen (1987) Christensen (1969) This study This study This study This study This study Christensen (1969) Christensen (1987) Simons & Vroman (1968, as t! walzi), Prud’homme van Reine (1970) This study This study This study This study This study This study This study This study This study Rousselet (1897) Christensen (1969) Davis & Gworek (1973)
LA = Louisiana,
NC = North
Carolina,
70
R.G. Verb, M.L. Vis, D.W. Ott & R.L. Wallace
through May 1998. Samples of macroscopic algae were collected and fixed in 2.5 % calcium carbonate-buffered glutaraldehyde. Maximum width, average depth, pH, specific conductance, temperature and mean current velocity were measured as described by Sheath et al. (1989). In addition, water samples were analyzed for orthophosphate, nitrate, total iron, aluminum and manganese with a DR/890 calorimeter, using standard protocols and powder pills from Hach Company (Anonymous, 1997). Water turbidity was measured with a Hach 21OOP’” turbidity meter. Species identifications of Kzucheria were determined by inducing gametophore production. Fresh material from the field was cultured in stream water for one week in a diurnal growth chamber at 15-20” C with fluorescent lighting providing 2OwOO lux under a 12-12 h photoregime. Species identification and nomenclature follows Rieth (1980) and Entwisle (1988). The rotifer was identified and nomenclature follows De Smet (1996). The length and width of galls containing~animals were measured using an 0~~1% micrometer. Images were taken with a Pixera digital camera attached to a BX40 Olympus microscope. A voucher specimen of the Kzucheria with galls has been deposited at the Ohio State University Herbarium (OS) under the accession number OS-399245. Collections
from North America
Kzucheria samples were collected in a number of locations from eastern North America and California (Tab. 1). These collections have been made sporadically over the past 30 years and some from the same location repeatedly. Sterile thalli with galls were transported back to the laboratory and placed in finger bowls with water from the collection site. The finger bowls were placed in a growth chamber at 12-15” C with fluorescent lighting providing 2000-4000 lux under a 14-10 hour photoregime. Cultures were left in the growth chamber for two weeks, after which time gametophore development had occurred and species could be identified. Unfortunately, no material was available for deposit in the Ohio State University Herbarium.
RESULTS
AND DISCUSSION
Seasonal Study Prior to September 1997 (May-August 1997), no visible Vuucheriu thalli had been collected from the Hocking River tributary location. In September 1997, Vaucheriu was found growing as a conspicuous mat in a large stream pool. Upon microscopic examination of Kzucheriu thalli, galls were observed (Fig. 1). The organism living in the galls was identified as the rotifer Proules werneckii. Vaucheriu with Proales wemeckii was present in stream collections from September 1997 through January 1998. Proales galls were most abundant in the October 1997 sample, but galls were present during all five months. Neither Kzucheriu nor associated Proales werneckii were collected in February 1998 or subsequent collections (March-May 1998). Therefore, it appears Proales wemeckii rapidly infects Vaucheriu mats since the first appearance of the thalli already shows gall formation. In addition, Proales is present throughout the tiucheriu growing season. Throughout the five sampling months, the Vaucheriu specimens collected were vegetative, lacking gametophores. Vaucheria specimens from the September sampling
Records of Vaucheria with Proales
71
were induced to form gametophores in culture and identified as two distinct species, Vaucheria geminata and K bursata (Figs 2, 3). Both species were colonized by Proales werneckii, possibly demonstrating lack of species specific infection. Measurements of width and length for 56 galls containing Proales wemeckii from the October sampling varied considerably. Galls ranged from 90-260 pm in width and 140-790 pm in length with a mean gall size of 135 x 320 pm. This wide range in gall dimensions may be due to progressive stages of gall development. Stream conditions were monitored throughout the sampling period (May 1997May 1998). The associated Kzucheria and Proales werneckii were able to tolerate a wide range in stream temperature, which fluctuated from 4 to 23” C. However, they are only found in association for a short period of time in the fall to early winter. During this time stream pH was neutral. Specific conductance was high (680-794 @cm-‘) for the region probably due to dissolved metals (Fe, Al and Mn). Orthophosphate levels were high and probably caused by a variety of anthropogenic inputs such as leachates from septic tanks and refuse present along the edge of the stream. These results suggest that Kzucheria bursata, V. geminuta and Proales wemeckii have a high tolerance of both organic and inorganic pollution. The period during which tiucheria with Proales was collected did not appear to have stream conditions uncharacteristic for the remainder of the sampling time.
Collections from North America Kzucheria species from California on the Pacific coast and Alabama, Georgia, Louisiana, North Carolina, Ohio and Tennessee in the eastern US were collected with Proales wemeckii galls (Tab. 1). These collections represent 13 new localities from states which, for the most part, had no previous records of this association. In North America, these associated taxa have only been reported from one Florida spring location (Davis & Gworek, 1973), California (as Debsalga gigasporangiu) (Habeeb, 1965) and noted to occur with no specific information on location (Wolle, 1887). Our collections confirm that this association is more prevalent in North America than previously thought. These taxa occurring together are probably cosmopolitan, but detailed information on geographical distribution suffers from the lack of widescale investigation. Worldwide, reports of this association have been sporadic with specific sites from Australia (Cribb, 1964; Christensen, 1987; Entwisle, 1988), Switzerland and France (Christensen, 1969), the United Kingdom (Rousselet, 1897) and the Netherlands (Prud’homme van Reine, 1972; Simons & Vroman, 1968) (Tab. 1). Again, when researchers have conducted extensive surveys of particular regions, Pro&s wemeckii in conjunction with Vaucheria species has been found in a number of localities (Christensen, 1969, 1987; Entwisle, 1988). Other authors have made reference to the association, in either literature reviews or early research, hut little information on location was included (Balbiani, 1878; Bock, 1954; Debray, 1890; Dormer, 1966; Ehrenberg, 1834; Hofker, 19 18; Gabriel, 1922; May, 1989; Rieth, 1980; Rothert, 1896; Weidner, 1952; Wolle, 1887). Although information is scarce, it would appear from the literature that this association has been known for quite some time. In this study, five distinct species were identified with Pro&es werneckii galls (Tab. 1). Vaucheria discoidea, V geminata and V pronu are new records of species forming galls and only K bursutu and K terrestris had been previously noted. There are at least 12 Vaucheriu species known worldwide to have associatedProules wemeckii including those reported in this study (Tab. 1). All Vaucheriu speciesreported forming galls are from sectionsComiculutue. Racemosue,T’buligerae, Kzucheria and Woroninia,
Records of Vaucheria with Proales
73
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