Final Report to the United States Fish and Wildlife Service
Report of The Status and Distribution of Ouachita Dusky Salamander, (Desmognathus brimleyorum) in the Ouachita Mountains of Oklahoma
Submitted by:
Doyle L. Crosswhite
and
Stanley F. Fox
Oklahoma Cooperative Fish and Wildlife Research Unit
and
Department of Zoology
Oklahoma State University
Stillwater, Oklahoma 74078
31 May 1995
LITERATURE REVIEW FOR THE OUACHITA DUSKY SALAMANDER,
Desmognathus brimleyorum
Stejneger (1895) first described the Ouachita Dusky Salamander, (Desmognathus brimleyorum) as a relatively large and robust plethodontid salamander that is often aquatic or semiaquatic. Although it is known that D. brimleyorum is restricted to the Ouachita Mountains (Conant and Collins 1991), there has been only limited study of the status of the species (Burt 1935; Grobman 1950; Means 1974). Until recently the range of this species was poorly defined and its abundance virtually unknown. Aside from this study, the most recent collections were made from 1977 and 1984 when Karlin et al. (1993) obtained 861 individuals from 49 locations, 11 of which were in Oklahoma.
The Ouachita dusky salamander generally inhabits areas near streams, springs and seeps where it is often found under submerged rocks, woody debris or buried in wet gravel (Conant and Collins 1991). The physiographic region which is home to this species extends from Atoka County, Oklahoma, east approximately 300 km to near Hot Springs, Arkansas (Dowling 1956). This narrow mountainous strip of unique pine-oak forest is only 80-90 km wide in places (Bruner 1931) and is made up of a series of east-west ridges and valleys that lie between the Gulf Coastal Plain to the south and the broad Arkansas River valley to the north (Fig. 1). Elevations range from 150 to 760 m (Dowling 1956). The positioning of the Ouachita Mountains effectively isolates the region from similar habitat found in the Ozark Plateau and the southern Appalachian Mountains. The relatively small island created by this topographical situation is home to several endemic salamander species as well as D. brimleyorum (Conant and Collins 1991). Other species include the Many-ribbed Salamander (Eurycea multiplicata), Rich Mountain salamander (Plethodon ouachitae), Caddo Mountain salamander (Plethodon caddoensis), and Fourche Mountain salamander (Plethodon fourchensis).
The Ouachitas are made up mostly of sedimentary rocks: sandstone, limestone, and conglomerate, along with metamorphic rocks such as shale and chert (Mohlenbrock 1993). Soils are predominately silty clay and silty loam and are very shallow and stony on the ridgetops, becoming progressively deeper downslope. These soils are of medium texture and are moderately permeable (Reagan 1974).
Several geologic factors contribute to the unique flora and fauna of the region. (1) Unlike the southwestern United States, the Ouachitas were not covered by shallow inland seas during the Cretaceous period and served as an island refuge for species. (2) The region also may have served as a refuge for plants and animals during the Pleistocene epoch, when glaciers covered adjacent northern regions. (3) During the late Cenozoic era, sediments that were deposited by inland seas were eroded, further defining boundaries and isolating the uplift. (4) Finally, during the Pleistocene, the river systems were formed (Dowling 1956).
Topography in the mountainous areas of the Ouachitas is often too steep for intensive agricultural use. This has led to a local economy which is heavily reliant upon livestock, poultry production, and a large timber industry.
Even-aged silviculture employing clearcutting, site preparation, and planting of pines has been the primary method of pine regeneration in these forests for the last 25 years. Although young pine plantations provide excellent habitat for many wildlife species adapted to early successional stages (such as deer, rabbits, and quail), even-aged management on short rotations, as typically practiced by the forest industry (especially on private lands, may be detrimental to those species that require mature forest habitat features (Thill 1990).
Clearcutting can cause changes in soil structure, hydrology, and both horizontal and vertical vegetation structure that subsequently affect temperature and moisture regimes. These altered characteristics in turn affect microhabitats that are important to salamanders (Ash 1988; Bury 1983; Heatwole and Lim 1961; Heatwole 1962; Matlack 1994; Pechman et al. 1991; Pough et al. 1987). Also, the water quality of streams may be degraded by increased sedimentation. These changes in microclimatic conditions on the forest floor and the erosion of stream quality are in part facilitated by canopy removal, elimination of the moisture-retaining forest floor litter, and soil compaction (Bratton 1994; Bury 1983; Raymond and Hardy 1991).
Contemporary logging practices have also altered the spatial and temporal disturbance regimes of forest ecosystems (Bratton 1994). Pough et al. (1987) suggested that small scale modifications to a forest may have little effect on salamander populations; after clearcutting, however, return of Plethodon cinereus was slow due to inadequate litter accumulation, which appeared to be a prerequisite for colonization. In general, deciduous leaf litter seems to be a very important habitat requirement for many terrestrial salamander species. Deciduous leaf litter retains moisture that plays a significant role in the distribution and activity patterns of terrestrial salamanders (Jaeger 1971). Pure stands of conifers are generally unsuitable for salamanders in the eastern and central United States (Bennett et al. 1980; Pough et al. 1987; Williams and Mullin 1987). In loblolly-shortleaf pine (Pinus taeda and P. echinata) stands of east Texas, Whiting et al. (1987) found that understory development and the degree of deciduous litter accumulation strongly influenced herpetofaunal communities. Petranka et al. (1993) compared clearcuts <5 years old with mature stands >80 years old and found that terrestrial salamanders were completely eliminated or reduced to very low numbers after the mature forest was cut. Petranka et al. estimated that 75-80% of salamanders from a variety of taxonomic groups are lost following timber harvest by clearcutting. Furthermore, it is estimated to require a century or more for populations to return to predisturbance levels following clearcutting (Petranka et al. 1994). There is concern that this reduction could produce population bottlenecks that result in decreased genetic diversity. In some cases local populations may be prone to extinction.
On a regional scale, survival of a reduced population depends upon recolonization through immigration from undisturbed areas (Fahrig and Merriam 1994). Constraints on such immigration, however, are that (1) salamanders generally only migrate under a narrow set of environmental conditions, (2) migrating individuals may have difficulty establishing territories in new areas due to interspecific competition with other herps, and (3) adult salamanders are often strongly philopatric (Petranka et al. 1993; Petranka 1994). As a result of these factors, recolonization of heavily disturbed areas is slow.
Salamanders are important components of the food chain and contribute a surprising amount of biomass to the community (Burton and Likens 1975; Pough et al. 1987). For example, population densities of Plethodon cinereus in the deciduous forests of the eastern United States have been recorded as high as 0.9-2.2 individuals/m2 (Heatwole 1962; Jaeger 1980). Furthermore, because salamanders are often habitat specialists with restricted distributions, they may be valuable indicator species to reveal the overall health and stability of the ecosystem. Despite the evidence that salamanders are important components in many ecosystems, these animals continue to be neglected by land managers (Pough et al. 1987). Some management plans may even promote mid-successional stages to maximize alpha diversity of other taxa at the cost of sensitive reptile and amphibian species (Faaborg 1980; Sampson and Knopf 1982). Recently, public awareness of the importance of the wildlife community as a whole has led to the concern for nongame wildlife and associated habitats (Jones 1986). One factor of concern regarding Desmognathus brimleyorum is the concordance of its range with an intensive timber industry. Several timber companies in the Ouachita Mountains practice clearcutting, which may potentially alter hydraulic features around springs and seeps that are primary habitats for the species. It is due to the species' limited distribution and potential threat with regard to habitat alteration that has stimulated interest in D. brimleyorum as a candidate for threatened status.
OBJECTIVES
The objective of this study was to determine the current status and distribution of the Ouachita Dusky Salamander (D. brimleyorum) by sampling localities of known historic occurrence and reviewing museum records to document any changes in distribution or abundance. This information is necessary to evaluate the impact, if any, that anthropogenic activities may have upon the species as well as to help guide future management decisions. Notes on microhabitat collected during the study add to our current knowledge of the biology of this species.
MATERIALS AND METHODS
We compiled regional collection records to evaluate the historical distribution of the Ouachita Dusky Salamander (Appendix I). We obtained records from the following sources: Oklahoma State University, University of Oklahoma, University of Kansas, University of Texas, American Museum of Natural History, and consultation with other researchers. During spring and fall 1993 and spring 1994, we visited a total of 55 sites in southeastern Oklahoma (Appendix II). Both sites of historic occurrence and those where the species could potentially occur were sampled. A representative survey of streams, springs and seeps was conducted in Choctaw, Latimer, LeFlore, McCurtain and Pushmataha counties in southeastern Oklahoma. The total number of sites visited by county were McCurtain, 20; LeFlore, 22; Latimer, 6; Pushmataha, 5; Choctaw, 2. Our sampling sites were selected based on historical distribution of D. brimleyorum and presence of available habitat as determined by Geographic Information System (GIS) analysis. Predicted areas of available habitat were identified after completing the 1993 field season, during which time we visited historic collection sites throughout southeastern Oklahoma. After confirming the presence of substantial populations of D. brimleyorum at several locations, we used GIS to identify appropriate conditions of habitat parameters known to be important to plethodontid salamanders. Assuming that the species attains its greatest densities in those locations with ideal habitats we were able to select a set of optimal habitat characteristics. The following habitat parameters were identified to be a subset of those believed to be important to the species: soil type, land use category, slope, aspect, and elevation. For each parameter an ideal range was determined within which we expected D. brimleyorum to be found. A set of maps was produced indicating the predicted range for the species in southeastern Oklahoma, i.e., those areas satisfying all the habitat criteria. After returning to the field to sample sites within the predicted range, we were able to identify additional extant populations. After collecting habitat information on these new sites we constructed a final set of maps that we believe accurately predicts the range of D. brimleyorum within the five county region surveyed.
Salamanders were sampled by dip net, 3.6 m x 1.8 m nylon seine (3.2 mm mesh), and by hand. Microhabitat parameters (e.g., stream depth and width, water temperature, and substrate type for aquatic locations; and slope, aspect, solar exposure, litter depth, substrate, and moisture for aquatic and terrestrial locations) were collected at each site where appropriate. Land-use practices were determined at these locations using GIS. Population density was estimated by determining the number of salamanders encountered during time-constrained searches.
Voucher specimens were taken in order to verify identification of larvae. Based upon estimates of density, salamanders were taken in a manner designed for minimal impact on the local population. Specimens were preserved in 10% formalin and transported to the laboratory where they were examined and transferred to a solution of 45% isopropyl alcohol. These reference collections were catalogued into the Collection of Vertebrates, Department of Zoology, Oklahoma State University.
RESULTS
Ninety one D. brimleyorum were observed at 16 localities in southeastern Oklahoma. We located extant populations at 4 localities in McCurtain county, 8 in LeFlore county, 1 in Latimer county, and 3 in Pushmataha county (Appendix II). Populations were most dense in LeFlore county along north-facing slopes of Winding Stair and Rich Mountains within the Ouachita National Forest.
We found D. brimleyorum and several other plethodontid salamanders, Southern Redback Salamander (Plethodon serratus), Northern Slimy Salamander (P. glutinosus), Rich Mountain Salamander (P. ouachitae), Sequoyah Slimy Salamander (P. sequoyah), Kiamichi Slimy Salamander (P. kiamichi), and Many-ribbed Salamander (Eurycea multiplicata) in riparian zones adjacent to and within high gradient streams of the Ouachita Mountains of southeastern Oklahoma. Desmognathus brimleyorum was most often found in or near 1st and 2nd order streams with depths ranging from 3-20 cm and widths from 0.2-5.0 m. Salamanders seemed to be associated with streams having a well developed deciduous canopy. Water temperatures at sites where D. brimleyorum were present ranged from 4-21o C. Structural heterogeneity of the stream habitat seemed to be important as well as slope and aspect. Dusky Salamanders were most often found in streams that contained coarse rocky or gravel substrates with at least some rocks >10 cm in width and where both pools and riffles existed. We never found D. brimleyorum in or near streams with sand or silt substrates. The salamanders utilized cracks and crevices in and between rocks for shelter. We found most adult salamanders under rocks in flowing water; most larvae were observed on the bottom of partially shaded pools. We also observed that salamanders typically did not occur in streams that supported large populations of predatory fish; e.g., sunfish, (Lepomis sp.); bass, (Micropterus sp.); and catfish, (Ictalurus sp.).
From the habitat conditions associated with all extant populations of D. brimleyorum, we refined our predictive habitat criteria (Table 1) and produced final maps of southeastern Oklahoma that predict the occurance of the species.
CONCLUSIONS AND RECOMMENDATIONS
In Oklahoma, the Ouachita Dusky Salamander appears to be locally abundant given the appropriate habitat conditions. Although the species may attain relatively high densities in and around seeps, springs, and high gradient streams, we believe the species to be sensitive to certain disturbances that could disrupt the hydrology of these fragile habitats. Consequently, we found the most productive sites within the Ouachita National Forest, protected from intensive agricultural practices. We found D. brimleyorum almost exclusively in upland areas and populations appear to be geographically isolated by expanses of lowland habitat and river flood plains. These lowland areas also tend to be centers for agriculture, which may further inhibit gene flow between populations.
There are, however, some limitations to this methodology. (1) The land use/cover categories produced by GIS are coarse. We believe, however, that the use of such ancillary information as soils and topography refine the predicted occurance so as to make the maps useful. (2) The habitat may be more extensive than the population. That is, even though the correct set of habitat conditions may exist at a site, geographic isolation of this relatively immobile species may result in the absence of a salamander population or the habitat patch may be too small or too fragmented to sustain a viable population. In order to overcome this problem one must increase the number of sample sites within the predicted range.
With respect to changes in the status and distribution of the species, we found the Ouachita Dusky Salamander to be absent from 8 of 18 historic collection sites examined. At least one new population was discovered that is noteworthy. We found an isolated population of D. brimleyorum in an isolated upland stream called Goat Creek (T3N,R20E,S3) in northern Latimer County. To the best of our knowledge, this is the western-most record for the species.
Dry conditions during the summer of 1993 and spring of 1994 prevented us from making an ideal systematic examination of the entire region within which D. brimleyorum is known to occur. Several of the sites we visited during this period seemed to have appropriate microhabitat characteristics, but stream flow was subterranean; salamanders are likely inactive under such conditions. With these considerations in mind, we make the following recommendations for future study of Desmognathus brimleyorum.
1. Study of microhabitat preferences and natural history of the Ouachita Dusky Salamander. Knowledge of microhabitat requirements and natural history is vital for management and protection of D. brimleyorum in Oklahoma, but only limited work has been done on these topics. We recommend a more rigorous examination of microhabitat preferences for streamside salamanders in order to determine their sensitivity to agricultural activities common in the region.
2. Assessment of the geographic pattern of genetic variation. We recommend that genetic diversity of isolated populations of D. brimleyorum be assessed using electrophoretic analysis to determine geographic patterns of genetic variation. Knowledge gained from this type of work would provide information for questions regarding management of the species. For example, can the species be treated as a single management unit, or should efforts be made to maintain several different populations? In fact, are all the populations really one species? Recent genetic work on the Slimy Salamander complex in this region (Highton et al. 1989) has identified 13 seperate species from what was originally considered one species (P. glutinosus). Management decisions may depend on the geographic distribution of genetically distinct populations.
Acknowledgements
The authors wish to thank David Leslie and the staff of the Oklahoma State University Cooperative Fish and Wildlife Research Unit, Oklahoma State University, and the Oklahoma Department of Wildlife Conservation for financial and technical support. We are grateful to Paul Shipman, Joy Yoshioka, Mindi Crosswhite, Mark Howery, Jason Evans, and Cory Fincher for their assistance with data collection. We thank Mark Gregory and the Oklahoma State University Department of Agriculture for assisting with GIS analysis.
APPENDIX I
Historic collection sites for the Ouachita Dusky Salamander, Desmognathus brimleyorum
Oklahoma: LeFlore Co.
1. Edge of Cucumber Cr., 1/8 m W of Jct. Hwy 259 (T1N,R25E,S7)
2. Winding Stair Mt., Hwy 1, 1.6-1.2 m W Jct. Hwy 259 (T3N,R25E,S21)
3. North slope of Winding Stair Mt., .2 m W Horse Thief Springs (T3N,R25E,S21)
4. North slope of Winding Stair Mt., below Horse Thief Springs (T3N,R25E,S7)
5. North slope of Rich Mt., 8.3 m W of the Arkansas state line on Hwy 1 (T2N,R25E,S1)
6. North slope of Rich Mt., 7.9 m W of the Arkansas state line on Hwy 1 (T2N,R25E,S1)
7. Round Mt. Forest Rd. 6025, 0.8 m NE Jct forest Rd. 6026 (T5N,R26E,S5)
8. Little Cedar Cr., 5.0 m E of Big Cedar (T2N,R25E,S7&8)
9. Kiamichi R. and tributaries, Hwy 63 ~7-9 m W of Arkansas state line (T2N,R26E,S18&29)
10. Hwy 270 E of Page (T3N,R27E,S20&30)
11. Hwy 1 near Jct. 259 (T3N,R25E,S20&26)
12. Hwy 1 near Buffalo Wallow Mt. and Jct. Hwy 271 (T4N,R23E,S19&23)
13. Wheeler Hollow Cr. (T4N,R24E,S16&20)
14. Beech Cr. (T1N,R26E,S11&27)
15. Hwy 63 (T2N,R26E,S20&24)
16. Cow Cr. Mt., (T1N,R27E,S20)
17. Murry Cr. (T1N,R27E,S28)
18. near Hwy 63 (T2N,R26E,S20&33)
19. Big Eagle Cr. (T1N,R24E,SE1/4,S3)
20. Blackfork Mt., W of Arkansas state line near Hwy 270 (T3N,R27E,S22)
21. East of Page (T3N,R26E,S24)
Oklahoma: Latimer Co.
1. Hwy 1/63 SW of Buffalo Mt., (T3N,R20E,S10&26)
2. Hwy 63 near Lake Carl Albert (T3N,R21E,S10)
3. Jackson Cr., near Hwy 63 (T3N,R21E,S21)
Oklahoma: McCurtain Co.
1. Two Mile Cr. near Hwy 70 (T6S,R27E,S5)
2. West of Smithville, Eagle Fork Hwy 259 (T1S,R25E,S22)
3. Mountain Fork River bridge Hwy 70 (T6S,R26E,S7)
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