and the cannibal snail ( Haplotrema con- 

 cavum ). 



Vertebrates 



Zone V shares vertebrate species coiri- 

 mon to uplands as well as Zone IV, The 

 large, spotted salamander ( Ambystona macu- 

 latum ) and mole salamander (A. t alpoideum ) 

 seem confined to Zone V. The red sala- 

 mander ( Pseudotriton ruber ) is shared with 

 Zone IV. Two common upland species are 

 the ubiquitous sMmy salamander ( Plethodon 

 qlutinosus ) and the red-backed salamander 

 ( Plethodo n cinereus ). Two toads, the nar- 

 rowmouth ( Gastrophryne carol inensis ) and 

 spadefoot ( Scaphiopus holbrooki ), Tnhabit 

 sandier portions of Zone V. In Zone V are 

 skinks of upland mesic slope forests, such 

 as the ground skink (Leiolopisma ) and 

 Eumeces inexpectatus , in addition to JE. 

 fasciatus . Among the snakes recorded are 

 the copperhead, canebrake rattlesnake 

 ( Cratalus horridus atricaudatus ), northern 

 brown fStoreria dekayi ), garter ( Thamno- 

 phis sirtal is ), rough green ( Opheodrys 

 ae stivus ), and ribbon snakes. Occasion- 

 ally, even upland species such as the 

 black racer and coachwhip ( Masticophis 

 flaqellum ), are found. We do not know how 

 many species migrate annually from upland 

 areas into Zone V when high water recedes, 

 or conversely, from Zone V to the uplands 

 during short periods of high water. 



In the Congaree Swamp the common 

 yellowthroat, pine warbler, wood thrush, 

 and eastern wood peewee seem to prefer 

 Zone V habitats. Zone V is perhaps the 

 preferred nesting and feeding ground of 

 the wild turkey (Figure 49). North Caro- 

 lina's only breeding colonies of cerulean 

 warblers (outside the Blue Ridge Moun- 

 tains) and Mississippi kites occur in a 

 60-km (37-mi) section of old growth timber 

 along the levees of the Roanoke River, 

 two-thirds of which (a 20C-m or 656-ft 

 wide strip) is dominantly Zone V vegeta- 

 tion. A number of birds that are commoner 

 in Zones IV and V than in other zones 

 include the white-breasted nuthatch, 

 Swainson's warbler, Carolina wren, and 

 yellow-throated vireo. Breeding bird 

 densities are generally higher in the 

 floodplain than in adjacent upland forests 

 (Dickson 1978). Kennedy (1977) noted that 

 more birds preferred Zone IV and V hard- 

 woods than other dominance types (e.g., 

 cottonwood-wil low-sycamore or cypress- 

 tupelo). 



Zones IV and V are the principcl 

 environments of the rare and endangered 

 ivory-billed woodpecker, Bachman's warbler 

 and probably the cougar (Wharton et al, 

 1981). Black bears (on Bear Island) con- 

 gregate on the higher, unlogged, acorn- 

 rich Zone IV and V bottomlands. Upland 

 forest forms sometimes occurring in Zone V 

 are the least shrew ( Cryptotis ), pine vole 

 ( Pitimys ). and, rarely, the common mole. 

 Other mammals are the same as reported for 

 Zone IV. 



Although Zone V environments may com- 

 prise a relatively small part of the total 

 floodplain acreage (for example, only 5% 

 in Congaree Swamp), these old levee 

 ridges are extremely important in the life 

 histories of many floodplain species. 

 They provide food, winter hibernacula, and 

 for the more terrestrial forms, high water 

 refuge and migration and dispersal routes. 

 In a number of southern swamps lacking a 

 Zone V, mounds of earth ("cattle mounts") 

 often were constructed by early human 

 residents to provide refuge for livestock 

 during high water. Tinkle (1959) found 

 narrow, long levees indispensible for the 

 egg-laying activities of many amphibious 

 snakes and turtles; he also discovered 

 that the swamp palm ( Sabal minor ) growing 

 there provided a major hibernaculum for 

 small vertebrates. 



THE USE 

 FISH 



OF BOTTOMLAND HARDWOOD ZONES BY 



Many fish species use Zones II 

 through V during inundation. At least 20 

 families and up to 53 species of fish 

 spawn and/or feed on the floodplain 

 (Lambou 1963; Holder et al. 1970, 1971; 

 Bryan et al. 1975, 1576; Huish and Pardue 

 1978; Walker 1980; Wharton et al. 1981; 

 and others). The catfish, sunfish, gar, 

 perch, and sucker families are particu- 

 larly well represented. 



Fish depend on an annual water level 

 fluctuation to limit intra- and interspe- 

 cific competition for food, space, and 

 spawning grounds (Lambou 1959). Fish dis- 

 tribution and abundance are thus keyed to 

 this cyclic phenomenon (Lambou 195S, 1962; 

 Bryan and Sabins 1975; Hern et al. 1980). 

 As most swamp-wise fishermen know, the 

 time and extent of overflow control the 

 size of the year classes of black bass and 

 sunfish (Lambou 1962). On the Danube 



92 



