Distribution of freshwater sponges and bryozoans in northwest Indiana.ABSTRACT. Current biogeographical bi·o·ge·og·ra·phy n. The study of the geographic distribution of organisms. bi  o·ge·og distributions of freshwater
sponges (Porifera) and bryozoans (Ectoprocta and Entoprocta) are poorly
known in Indiana. Although seemingly ubiquitous in many aquatic
communities, neither group has received much notoriety in this state,
nor the midwestern United States United States, officially United States of America, republic (2005 est. pop. 295,734,000), 3,539,227 sq mi (9,166,598 sq km), North America. The United States is the world's third largest country in population and the fourth largest country in area. . In an initial effort to begin the
systematic taxonomic distribution of these groups in Indiana, this study
identified 2 sponges and 13 bryozoans in northwest Indiana Northwest Indiana, also known as The Calumet Region, or just The Region, is comprised of Lake, Porter, LaPorte, Newton, and Jasper counties in Indiana. This region neighbors Chicago, Illinois and Lake Michigan, and is also the Indiana component of the Chicago lakes. The
recent appearance of the zebra mussel zebra musselEither of two species of tiny mussels (genus Dreissena) that are prominent freshwater pests. They proliferate quickly and adhere in great numbers to virtually any surface. (Dreissena polymorpha Noun 1. Dreissena polymorpha - inch long mollusk imported accidentally from Europe; clogs utility inlet pipes and feeds on edible freshwater mussels zebra mussel ) in the Great Lakes Great Lakes, group of five freshwater lakes, central North America, creating a natural border between the United States and Canada and forming the largest body of freshwater in the world, with a combined surface area of c.95,000 sq mi (246,050 sq km). has focused new attention on the sessile sessile /ses·sile/ (ses´il) attached by a broad base, as opposed to being pedunculated or stalked. ses·sile adj. Permanently attached or fixed; not free-moving. benthic ben·thos n. 1. The collection of organisms living on or in sea or lake bottoms. 2. The bottom of a sea or lake. [Greek. communities, as significant changes in benthic community structure are expected with the introduction of this exotic species. Without knowing the historical and current distribution of native invertebrates, such as the sponges and bryozoans, it is not possible to identity changes in community composition over time. Keywords: Porifera, Ectoprocta, Entoprocta, freshwater bryozoan bryozoan Aquatic invertebrate of the phylum Bryozoa (“moss animals”), members (called zooids) of which form colonies. Each zooid is a complete and fully organized animal. Species range in size from a one-zooid “colony” small enough (less than 0. , freshwater sponge The distribution and taxonomy of freshwater sponges (Porifera: Spongillidae) in North America North America, third largest continent (1990 est. pop. 365,000,000), c.9,400,000 sq mi (24,346,000 sq km), the northern of the two continents of the Western Hemisphere. has not received widespread attention, although some notable works exist for the entire United States (Penney & Racek 1968), and regions surrounding the Great Lakes (Potts 1887; Smith 1921; Old 1931; Jewell 1935; Neidhoefer 1940; Eshleman 1950; Ricciardi & Reiswig 1993). Ricciardi & Reiswig (1993) found 15 different species and suggested that additional taxa taxa: see taxon. could be identified in eastern Canada Eastern Canada (also the Eastern provinces) is the region of Canada generally considered to be east of Manitoba, consisting of the following provinces:
n. The biological study of the geographic distribution of animals, especially the causes and effects of such distribution. zo distribution, but rather a lack of observation (Frost 1991). We have only been able to find six published records of sponge distributions in Indiana (Evermann & Clark 1920; Kintner 1938; Early et al. 1996; Lauer & Spacie 1996; Early & Glonek 1999; Lauer et al. 2001), and these are limited in scope. Although bryozoans (Ectoprocta and Entoprocta) are widely distributed Adj. 1. widely distributed - growing or occurring in many parts of the world; "a cosmopolitan herb"; "cosmopolitan in distribution" cosmopolitan bionomics, environmental science, ecology - the branch of biology concerned with the relations between organisms in epibenthic and littoral littoral /lit·to·ral/ (lit´ah-r'l) pertaining to the shore of a large body of water. littoral pertaining to the shore. communities (Rogick 1934, 1957; Bushnell et al. 1987; Ricciardi & Lewis 1991), little is known about their zoogeo-graphical status. There are 25 species known in North American North American named after North America. North American blastomycosis see North American blastomycosis. North American cattle tick see boophilusannulatus. freshwater (Wood 2001; Smith 1992), and the distribution of many of these species is thought to be ubiquitous (Bushnell 1974). In the Great Lakes region The Great Lakes region can refer to:
taxon, taxonomic category Adapid, Adapid group - extinct small mostly diurnal lower primates that fed on leaves and fruit; abundant in North America and Europe 30 to 50 million years in Indiana except for these recent studies in the northern part of the state (Wood 1996; Lauer et al. 1999; Last & Whitman 1999/2000). Without knowing the historical and current distribution of native sponges and bryozoans, it is not possible to identify changes in community composition over time. Rapid modification in the quantity and diversity of benthic species has been observed in the Great Lakes with the appearance of non-indigenous species, such as the zebra mussel (Dreissena polymorpha). These invasions have threatened the biotic biotic /bi·ot·ic/ (bi-ot´ik) 1. pertaining to life or living matter. 2. pertaining to the biota. bi·ot·ic adj. 1. Relating to life or living organisms. integrity of native organisms and negatively affected the ecosystem stability of sessile benthic communities (Hebert et al. 1991; Mills et al. 1994). The objectives of this study were to determine the biogeographical distribution of freshwater sponges and bryozoans in the lakes of Lake, Porter and LaPorte counties, Indiana (excluding Lake Michigan), and to identify the water quality habitats of both bryozoans and freshwater sponges where they are found. METHODS Twenty lakes in three northwest Indiana counties were sampled for freshwater sponges and bryozoans during the summer of 1998 (Fig. 1). The lakes sampled were public, freshwater natural lakes remnant from the geological activity that occurred during the Pleistocene glaciation The Pleistocene glaciation, also known as the Pleistocene ice age or simply the ice age, was one of the most significant events in recent Earth history, occurring in the Pleistocene epoch (1,808,000 to 11,550 years ago). approximately 12,000 years ago (reviewed by Hutchinson 1957). The size and maximum depths of the lakes varied (Table 1). In each lake, a single collection effort occurred in June-July and typically included 2-4 man-hours of shallow water See:
Swimming done underwater with a self-contained underwater-breathing apparatus (scuba), as opposed to skin diving, which requires only a snorkel, goggles, and flippers. Scuba gear was invented by Jacques-Yves Cousteau and Émile Gagnan in 1943. was used in deeper waters when warranted. Sticks, plants, rocks, and other similar hard substrate were examined for the sponges and bryozoans. Collections entailed picking or scraping the organisms from the substrate, or removing the organism with the substrate still attached. Sponge identification is based largely on spicule spicule: see chromosphere. morphology and care was taken to collect specimens bearing gemmules This article is about the mechanism for heredity. For the internal buds of freshwater sponges, see Gemmule. Gemmules were imagined particles of inheritance proposed by Charles Darwin as part of his Pangenesis theory. , as well as somatic somatic /so·mat·ic/ (so-mat´ik) 1. pertaining to or characteristic of the soma or body. 2. pertaining to the body wall in contrast to the viscera. so·mat·ic adj. tissue. Similarly, bryozoan identification typically requires the inspection of statoblasts in addition to the vegetative vegetative /veg·e·ta·tive/ (vej?e-ta?tiv) 1. of, pertaining to, or characteristic of plants. 2. concerned with growth and nutrition, as opposed to reproduction. 3. colony, and both were collected when possible. Most species are large enough to observe with the naked eye, but a hand lens hand lens n. A hand-held magnifying glass. was used to validate colonies that were small or non-descript. Samples were preserved immediately upon collection with 70% ethyl alcohol ethyl alcohol: see ethanol. , and returned to the laboratory for taxonomic identification. Laboratory preparation of sponge spicules (megascleres, microscleres, and gemmoscleres) was performed using an acid digestion, followed by washings in water and ethyl alcohol. The alcohol was evaporated, and the spicules were mounted in Permount. This technique creates a permanent mount and allows the microscopic examination of the spicules as described in Pennak (1989). Species were identified using the taxonomic keys of Pennak (1989), Frost (1991), and Ricciardi & Reiswig (1993). Bryozoan colonies and statoblasts were examined using a binocular microscope binocular microscope n. A microscope having two eyepieces, one for each eye, so that the object can be viewed with both eyes. at 10X magnification and identified using the taxonomic keys of Pennak (1989), Wood (1989, 2001), Ricciardi & Reiswig (1994) and Wood (1996). Water quality measurements were taken at each location at the time of biological sampling to characterize the habitat where bryozoans and sponges were found. Physical and chemical parameters measured were: temperature, pH, Secchi, conductivity, hardness, calcium, magnesium, total dissolved solids Total dissolved solids (often abbreviated TDS) is an expression for the combined content of all inorganic and organic substances contained in a liquid which are present in a molecular, ionized or micro-granular (colloidal sol) suspended form. , and dissolved oxygen. Chemistry analysis was completed within 6 h of collection using Hach Company methodology (Hach Chemical Company 1988, 1990) for oxygen (DO), hardness, calcium, magnesium, and total dissolved solids (TDS TDS total dissolved solids. ). A digital Marine Systems meter was used for pH measurements, while conductivity and temperature were taken on site using a Hach Model 44600 conductivity meter. RESULTS Thirteen bryozoans (3 classes, 7 families) were identified in the 20 study lakes (Table 2) and represent the first published account of these species from Indiana. Twelve of them are in the Phylum Ectoprocta Noun 1. phylum Ectoprocta - coextensive with or a subphylum of Bryozoa Ectoprocta Bryozoa, phylum Bryozoa, polyzoa - marine or freshwater animals that form colonies of zooids and follow the classification scheme outlined by Wood (2001), while a single species, Urnatella gracilis, is in the Phylum Entoprocta Noun 1. phylum Entoprocta - sometimes considered a subphylum of Bryozoa Endoprocta, Entoprocta Bryozoa, phylum Bryozoa, polyzoa - marine or freshwater animals that form colonies of zooids . Two freshwater sponge species, Eunapius fragilis and Spongilla aspinosa, were also found. Eunapius fragilis has been found previously in the Indiana waters of Lake Michigan (Lauer & Spacie 1996), while Spongilla aspinosa is a new record for Indiana. Physical and chemical measurements (Table 1) provided some reference to the environmental limits of the bryozoans and sponges. Although the samples for measurement were collected at the 1 m depth, they provided some indication of the range of tolerance of these species. This is particularly true of some species, such as Cristatella mucedo, that were found at multiple sites. DISCUSSION Finding 12 ectoproct bryozoan species (2 classes, 6 families) in a relatively limited three-county area in northwest Indiana was unexpected. As a comparison, Bushnell (1965a) sampled 122 sites in 48 counties in Michigan The boundaries of counties in the U.S. state of Michigan have not changed since 1897. However, throughout the 19th century, the state legislature frequently adjusted county boundaries. over four years and found 13 species, while Wood (1989) found 13 species from 60 locations in Ohio, and Ricciardi & Reiswig (1994) found 14 species from 80 localities in eastern Canada. Other investigators found similar numbers of species (Lake Erie Lake Erie Great Lake; once so polluted, referred to as Lake Eerie. [Am. Hist.: NCE, 887] See : Filth , Rogick (1934; Ottawa River Ottawa River River, eastern central Canada, the chief tributary of the St. Lawrence River. It rises in the Laurentian plateau of western Quebec and flows west to form the Quebec-Ontario border before joining the St. Lawrence west of Montreal. , Ricciardi & Lewis (1991); Louisiana, Everitt (1975)), although some recent changes in ectoproct classifications (see Wood 1996, 2001) may alter the number of species. Our findings suggest that the 20 lakes surveyed in Lake, Porter and LaPorte Counties have a diverse environmental character and support a wide variety of ectoproct bryozoan species despite the small biogeographical area surveyed. The most common ectoproct species by distribution were Fredericella indica, Plumatella reticulata, Pectinatella magnifica, and Cristatella mucedo (Table 2). The first three species are widely distributed in Ohio (Wood 1989) and Michigan (Bushnell 1965a), while C. mucedo is found throughout North America, but less frequently (Bushnell 1965a; Smith 1985; Wood 1989; Ricciardi & Reiswig 1994). Pectinatella magnifica colonies can grow to a large size (> 50 cm) and are readily found as a gelatinous gelatinous /ge·lat·i·nous/ (je-lat´i-nus) like jelly or softened gelatin. ge·lat·i·nous adj. 1. Of, relating to, or containing gelatin. 2. Resembling gelatin; viscous. matrix, particularly late in the summer. Although our specimens were not this large (10-30 cm), they were readily seen during collections. Wood (1989) states that this species also has a wide tolerance for water quality characteristics, but appears to be thermophilic ther·mo·phil·ic adj. Requiring high temperatures for normal development, as certain bacteria. , as growth of statoblasts into colonies is limited below 20[degrees]C. Water temperatures at the time of our collections in all 20 lakes were above this limit and did not appear to be inhibiting growth of this species. Fredericella indica, Plumatella casmiana, Plumatella emarginata, Plumatella repens and Paludicella articulata are widely distributed in the midwestern United States and North America (Brown 1933; Bushnell 1965a; Wood 1989; Ricciardi & Lewis 1991; Ricciardi & Reiswig 1994); and their occurrences in northwest Indiana were not unexpected. Despite the apparent widespread distribution throughout the U.S. (Wood 2001), Urnatella gracilis was found only in Cedar Lake Cedar Lake may refer to: Cities, towns, townships etc.
United States Two species of sponges, Eunapius fragilis and Spongilla nr. aspinosa, were found in 5 of the 20 lakes (Table 2). Eunapius fragilis is a common species in the midwestern U.S. (Old 1931; Jewell 1935; Lauer & Spacie 1996). It has been found on all continents and climates and is described as truly cosmopolitan by Harrison (1974). As might be expected, this species exhibits a wide tolerance to environmental conditions (summarized by Harrison 1974), well within the water quality measurements we observed (Table 1). The sponge has a preference for growing in areas where the canal system is not clogged by siltation (Potts 1887), with growth typically being enhanced on the underside of hard substrates (Lauer & Spacie 1996). Spongilla nr. aspinosa is far less common than E. fragilis and has been reported only from waters that have low pH (Potts 1887; Eshleman 1950; Ricciardi & Reiswig 1993). Identification of S. nr. aspinosa was tentative, as gemmules are required for positive confirmation; and we could not find any. Gemmules are over-wintering structures and typically used as a protection against adverse conditions. However, S. nr. aspinosa is environmentally tolerant and rarely produces gemmules as part of its life history strategy (Potts 1887). Although some question in identification remains for this species, this is probably the correct name for this organism. Associating water quality parameters with specific organism species was a bit difficult from our results with the limited sampling effort of this survey. However, these lakes are generally characterized as being eutrophic eu·troph·ic adj. Relating to, characterized by, or promoting eutrophia. , with extensive anthropogenic an·thro·po·gen·ic adj. 1. Of or relating to anthropogenesis. 2. Caused by humans: anthropogenic degradation of the environment. inputs. Both the sponges and the bryozoans in this study represent new records for Indiana. These records should not be considered range extensions or changes in habitats where they were found, but rather, simply an initial assessment of the sponge and bryozoan biogeographical distributions in Indiana. Many of these species are found throughout the geographical region, and might be expected. Considering the relative ease with which these organisms can be found, the paucity of studies on these organisms in Indiana or even nationwide does not suggest they have a limited distribution, but rather that they have not been studied. Our limited mid-summer sampling effort may have also missed some species that are present due to seasonal fluctuations or variations in the growth and development of these organisms. More intense and complete works on the subject, e.g., Wood (1989) and Ricciardi & Reiswig (1993, 1994), support this hypothesis. This study identified the biogeographical distribution of these little known groups in northwest Indiana, and by doing so, initialized an effort to more fully understand the significance and ecological importance of sponges and bryozoans to this region.
Table 1.--Lake basin morphometry, and physical and chemical measurements
of the water found at each of the 20 sampling locations during 1998.
Water samples were collected at the time of sponge and bryozoan
sampling.
Max. Max.
size depth
Country/Lake Date (ha) (m)
Lake
1. Cedar 7/10 316 5.0
2. Francher 7/10 4.0 12
3. Holiday 7/6 77 5.5
4. Lakeland 7/8 2.8 10
5. Lemon 7/10 13 4.1
6. Wolf 7/13 156 2.5
LaPorte
7. Clear 6/22 43 3.5
8. Pine 6/24 228 16
9. Saugany 7/3 30 20
10. Stone 6/23 51 11
Porter
11. Big Bass 7/6 28 3.4
12. Eliza 7/15 18 14
13. Flint 6/29 36 20
14. Long 6/30 26 8.2
15. Loomis 6/30 25 17
16. Louise 7/16 92 7.6
17. Minnehaha 7/13 8.9 11
18. Round 6/29 0.7 5.5
19. Spectacle 6/30 12 3.0
20. Wahob 6/29 8.0 15
Water quality parameter
Temp pH Secchi
[degrees]C S.U. m
County/Lake
Lake
1. Cedar 8.7 4.0
2. Francher 27.3 8.6 1.0
3. Holiday 31.0 7.8 0.3
4. Lakeland 27.3 8.7 4.0
5. Lemon 27.3 7.3 0.2
6. Wolf 31.6 8.8 1.8
LaPorte
7. Clear 28.9 9.4 1.0
8. Pine 29.8 9.0 2.0
9. Saugany 28.4 8.5 5.0
10. Stone 28.4 8.7 3.0
Porter
11. Big Bass 28.1 7.8 0.3
12. Eliza 28.0 9.4 0.4
13. Flint 29.4 8.7 1.5
14. Long 26.9 7.8 1.0
15. Loomis 29.1 8.4 0.5
16. Louise 28.2 8.4 1.6
17. Minnehaha 29.6 8.6 5.0
18. Round 29.1 7.5 2.1
19. Spectacle 29.5 8.5 1.4
20. Wahob 28.8 7.5 3.0
Water quality parameter
D.O. Hard. Ca
mg/l mg/l mg/l
County/Lake
Lake
1. Cedar 7.8 74 29.6
2. Francher 8.2 78 31.2
3. Holiday 6.9 76 30.4
4. Lakeland 7.8 74 29.6
5. Lemon 6.5 29 11.6
6. Wolf 8.0 65 26.0
LaPorte
7. Clear 9.0 60 24.0
8. Pine 9.3 39 15.6
9. Saugany 7.5 47 18.8
10. Stone 8.0 54 21.6
Porter
11. Big Bass 4.0 71 28.4
12. Eliza 10.7 61 24.4
13. Flint 7.0 67 26.8
14. Long 5.8 27 10.8
15. Loomis 5.8 53 21.2
16. Louise 7.5 75 30.0
17. Minnehaha 8.3 78 31.2
18. Round 6.0 41 16.4
19. Spectacle 5.8 57 22.8
20. Wahob 6.1 44 17.6
Water quality parameter
Mg Cond. TDS
mg/l [micro]mhos/cm mg/l
County/Lake
Lake
1. Cedar 10.8 911 456
2. Francher 11.4 386 193
3. Holiday 11.1 394 198
4. Lakeland 10.8 911 456
5. Lemon 4.2 416 212
6. Wolf 9.5 289 147
LaPorte
7. Clear 8.8 653 331
8. Pine 5.7 293 148
9. Saugany 7.0 432 218
10. Stone 7.9 259 136
Porter
11. Big Bass 10.4 338 170
12. Eliza 8.9 267 134
13. Flint 9.8 351 175
14. Long 4.0 221 111
15. Loomis 7.8 361 181
16. Louise 11.0 334 168
17. Minnehaha 11.4 550 276
18. Round 6.0 352 219
19. Spectacle 8.4 412 201
20. Wahob 6.4 271 136
Table 2.--Sponges and bryozoans found in Lake, Porter, and LaPorte
countries, Indiana. The lake site
number corresponds to the corresponding lake number in Figure 1.
Lake number
Bryozoans
Phylum Ectoprocta
Class Phylactomaemata
Family Fredericellidae
Fredericella indica Annandale 1909 2, 3, 6, 17
Fredericella browni (Rogick 1945) 8, 10
Family Plumatellidae
Hyalinella punctata (Hancock 1850) 2, 17
Plumatella casmiana (Oka 1907) 3
Plumatella emarginata Allman 1844 3
Plumatella fungosa (Pallas 1768) 1
Plumatella orbisperma (Kellicott 1882) 9
Plumatella nitens Wood 1996 1, 6, 12
Plumatella reticulata Wood 1988 1, 2, 3, 8, 10, 12, 13, 16
Family Pectinatellidae
Pectinatella magnifica (Leidy 1851) 3, 11, 12, 18, 20
Family Cristatellidae
Cristatella mucedo Cuvier, 1798 7, 8, 9, 10, 13, 16, 18, 20
Class Gymnolaemata
Family Paludicellidae
Paludicella articulata (Ehrenberg 1831) 13, 17
Phylum Entoprocta
Family Urnatellidae
Urnatella gracilis Leidy 1851 1
Sponges
Phylum Porifera
Family Spongillidae
Eunapius fragilis Leidy 1851 3, 13, 20
Spongilla nr. aspinosa Potts 1880 16, 18,20
ACKNOWLEDGMENTS We wish to thank the Indiana Academy of Science and Ball State University for funding. We also wish to thank Anthony Ricciardi, University of Guelph The University of Guelph is a medium-sized university located in Guelph, Ontario, established in 1964. While the U of G offers degrees in many different disciplines, the university is best known for its focus on life sciences, based in part on a long-standing history of , for taxonomic confirmation of the sponges and Timothy Wood For the figure skater, see . Timothy John Rogerson Wood, known as Tim Wood, (born August 13 1940) is a British politician. He was the Conservative Party Member of Parliament for Stevenage, which he won at the 1983 general election. , Wright State University, for his assistance with the bryozoan species identifications and review of the manuscript. We also thank Joseph Beatty for suggesting improvements in an early draft, Nicole M. Nuzzo, Amanda Reichler (diving and collecting assistants), Mitchell Alix (assistance with lake data), and the land owners who allowed us access to the lakes from their properties. LITERATURE CITED Brown, J.D. 1933. A limnological lim·nol·o·gy n. The scientific study of the life and phenomena of fresh water, especially lakes and ponds. [Greek limn study of certain fresh-water Polyzoa with special reference to their statoblasts. Transactions of the American Microscopical Society 52:271-313. Bushnell, J.H. 1965a. On the taxonomy and distribution of freshwater Ectoprocta in Michigan. Part 1. Transactions of the American Microscopical Society 84:231-244. Bushnell, J.H. 1965b. On the taxonomy and distribution of freshwater Ectoprocta in Michigan. Part 2. Transactions of the American Microscopical Society 84:339-358. Bushnell, J.H. 1965c. On the taxonomy and distribution of freshwater Ectoprocta in Michigan. Part 3. Transactions of the American Microscopical Society 84:529-548. Bushnell, J.H. 1974. Bryozoa (Ectoprocta). Pp. 157-194, In Pollution Ecology of Freshwater Invertebrates. (C.W. Hart & S.L.H. Fuller, eds.). Academic Press, New York New York, state, United States New York, Middle Atlantic state of the United States. It is bordered by Vermont, Massachusetts, Connecticut, and the Atlantic Ocean (E), New Jersey and Pennsylvania (S), Lakes Erie and Ontario and the Canadian province of . Bushnell, J.H., S.Q. Foster & B.M. Wahle. 1987. Annotated inventory of invertebrate invertebrate (ĭn'vûr`təbrət, –brāt'), any animal lacking a backbone. The invertebrates include the tunicates and lancelets of phylum Chordata, as well as all animal phyla other than Chordata. populations in an alpine lake Alpine Lake is a lake in Marin County, California.[1] Notes 1. ^ Topozone map, USGS, September 17, 2007
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