Publications (WR) Water Resources 3-1989 Red swamp crayfish ecology in Lake Mead Suzanne E. Leavitt University of Nevada, Las Vegas Jennifer Stephens Haley University of Nevada, Las Vegas Mikell Hager University of Nevada, Las Vegas Donald H. Baepler University of Nevada, Las Vegas Nevada Department of Wildlife Follow this and additional works at: http://digitalscholarship.unlv.edu/water_pubs Part of the Aquaculture and Fisheries Commons, Biology Commons, Environmental Monitoring Commons, Fresh Water Studies Commons, Natural Resources Management and Policy Commons, Terrestrial and Aquatic Ecology Commons, and the Water Resource Management Commons Repository Citation Leavitt, S. E., Haley, J. S., Hager, M., Baepler, D. H., Nevada Department of Wildlife (1989). Red swamp crayfish ecology in Lake Mead. Available at: http://digitalscholarship.unlv.edu/water_pubs/96 This Technical Report is brought to you for free and open access by the Water Resources at Digital Scholarship@UNLV. It has been accepted for inclusion in Publications (WR) by an authorized administrator of Digital Scholarship@UNLV. For more information, please contact [email protected]. C.I Environmental Research Center University of Nevada, Las Vegas Las Vegas. Nevada 89154 (702) 739-3382 BUREAU Or RECLAMATION DCNVER LIBRARY RED SWAMP CRAYFISH ECOLOGY IN LAKE MEAD Suzanne E. Leavitt Jennifer Stephens Haley Mikell Hager Dr. Donald H. Baepler DATE DUE SEP 23 (1997 Lake Mead Limnological Research C Environmental Research Center University of Nevada-Las Vegas In Cooperation With: The Nevada Department of Wildlif.f David A. Buck John Hutchings QAVLORO March 1989 Final Report PRINTED IN U S A ACKNOWLEDGMENTS This study was funded by the Nevada Department of Wildlife. We would like to thank them for their support. We would especially like to thank Dave Buck and John Hutchings of the Nevada Department of Wildlife for their cooperation and involvement in this study. In addition, we would like to thank Michele Salas for her help in the field and with the report graphics, typing, and editing. 11 ABSTRACT Red swamp crayfish (Procambarus clarkii) were trapped in Flamingo Wash, an urban wash of Las Vegas, during four periods of 1988. Life history and reproductive success were determined. The trapped crayfish were marked and released into a study cove in Lake Mead as part of an experimental stocking program. Retrap data from the study cove were used to determine life history, habitat preferences, and movement patterns of the stocked crayfish. In addition, a literature search was done on red swamp crayfish ecology, including food preferences, life history, habitat preferences and fish predation. ui ABSTRACT Red swamp crayfish (Procambarus clarkii) were trapped in Flamingo Wash, an urban wash of Las Vegas, during four periods of 1988. Life history and reproductive success were determined. The trapped crayfish were marked and released into a study cove in Lake Mead as part of an experimental stocking program. Retrap data from the study cove were used to determine life history, habitat preferences, and movement patterns of the stocked crayfish. In addition, a literature search was done on red swamp crayfish ecology, including food preferences, life history, habitat preferences and fish predation. ui Table of Contents Page I. Introduction 1 A. B. C. D. E. 1 3 4 6 6 History of Fisheries in Lake Mead Habitat Preferences Life History of the Red Swamp Crayfish Feeding Habits Predation on Crayfish II. Description of the Study Area IE. Methods 10 A. B. C. 10 10 D. E. F. IV. 7 Literature Review Lake-wide Crayfish Population Sampling Population Sampling and Life History Evaluation in Flamingo Wash Lake Mead Stocking Program 1. Marking and Release 2. Monitoring Predation by Game Fish Habitat Improvement by Vegetation Introduction Results A. B. C. D. E. F. 13 14 14 16 18 18 19 Literature Review Lake-wide Crayfish Population Sampling Population Sampling and Life History Evaluation in Flamingo Wash Lake Mead Stocking Program 1. Life History 2. Habitat Preferences 3. Movement Patterns Predation by Game Fish Habitat Improvement by Vegetation Introduction 19 19 19 22 24 24 27 45 47 V. Conclusions 47 VI. Literature Cited 48 IV List of Figures and Tables Page 8 Figure 1. Map of Lake Mead Figure 2. Lake Mead water elevations (in feet) from January 1984 through December 1988 9 Locations of red swamp crayfish sampling sites monitored between October 1986 and July 1987 11 Distribution of red swamp crayfish in Lake Mead from trap catches (October 1986 - July 1987) 12 Figure 5. Location of study cove 15 Figure 6. Trapline placement in the study cove 17 Figure 7. Percent of red swamp crayfish trapped in each sex/age class in Flamingo Wash during four trapping periods 21 Percent of red swamp crayfish trapped in each sex/age class in the study cove during four trapping periods 25 Number of red swamp crayfish trapped at various distances from the Lot 1, Lot 2, and Lot 3 release sites during the first stocking period 28 Figure 10. Number of red swamp crayfish trapped at various distances from the Lot 1, Lot 2, and Lot 3 release sites during the second stocking period 32 Figure 11. Number of red swamp crayfish trapped at various distances from the Lot 1, Lot 2, and Lot 3 release sites during the third stocking period 36 Figure 12. Number of red swamp crayfish trapped at various distances from the Lot 1, Lot 2, and Lot 3 release sites during the fourth stocking period 42 Table 1. Number of red swamp crayfish recaptured after each release 23 Table 2. Percent recaptured from previous releases 23 Table 3. Catch per unit effort of red swamp crayfish trapped in various habitats 26 Mean temperature (C), dissolved oxygen (DO - mg/1), conductivity (EC - micro mho/cm) and pH from the surface of the water to the bottom of the cove at the study site 46 Figure 3. Figure 4. Figure 8. Figure 9. Table 4. List of Appendices Appendix A. Annotated bibliography to crayfish ecology. Appendix B. Habitat sampled by each trap during each introduction period. VI I. INTRODUCTION A. History of Fisheries in Lake Mead The sport fisheries in Lake Mead are comprised mainly of largemouth bass (Micropteris salmoides), striped bass (Morone saxatilis), and rainbow trout (Salmo gairdnerii). Largemouth bass were introduced into the reservoir in 1935 shortly after the completion of Hoover Dam, and threadfin shad (Dorosoma petenense) were introduced in 1954 to expand the forage base for largemouth bass (Allan and Roden, 1978). The largemouth bass fishery began declining in 1963 after Glen Canyon Dam was constructed 456 km upstream. The largemouth fishery declined even more in the late 1970's, and the population is still severely depleted. Striped bass and rainbow trout were introduced in 1969 to augment the failing largemouth bass fishery. Although the lake has supported a substantial fishery in the past (Allan and Roden, 1978), current evaluations of the fishery have provided little hope for the future. The trout fishery collapsed in 1977, and the striped bass fishery began experiencing problems in 1978 when emaciated fish appeared in the population. Striped bass catch decreased drastically in 1979, and the population was comprised of small and often emaciated fish during the early 1980's (Paulson and Baker, 1984). The red swamp crayfish is an important food source for two prominent game fish in Lake Mead, the largemouth bass (Micropterus salmoides) and the striped bass (Morone saxatilis), as determined by creel census data collected by the Nevada Department of Wildlife (NDOW). Therefore, crayfish serve not only as an important link in the food chain of fish, but also play an important role in recycling nutrients from the benthos upwards into the higher trophic levels. The red swamp crayfish was first reported from lakes Mead and Mojave by Allan and Roden (1978). Jonez and Sumner (1954) had previously reported the introduction of Astacus sp. in the Willow Beach area of Lake Mohave by bait releases, but these records were not documented with collections. Two lake-wide crayfish surveys conducted in October 1986 and April 1987 indicated that population densities of P. clarkii in Lake Mead are very low. This species has, however, become established throughout the lake and is reproductively successful (Peck et al., 1987). These results are inconsistent with food habitat analysis completed by Albert and Baker (1982) which indicate that crayfish are being utilized heavily as forage during certain times of the year. This study is the result of a contract between the Nevada Department of Wildlife (NDOW) and the Lake Mead Limnological Research Center (LMLRC) to evaluate the potential for stocking red swamp crayfish (Procambarus clarkii) in Lake Mead and whether these crayfish will become reproductively successful resulting in a broader forage base for game fish. The objectives of this study were to: 1. Determine the distribution and relative abundance of red swamp crayfish in Lake Mead on a seasonal basis. 2. Determine the breeding season, reproductive success, growth patterns, and food preferences for red swamp crayfish in Lake Mead. 3. Study the life history of red swamp crayfish in Las Vegas urban washes to determine breeding season, reproductive success, and growth patterns. 4. Analyze patterns of crayfish predation. 5. Monitor the success of the vegetation transplant project and recommend specific plants that might be important to crayfish based on life history and microhabitat data collected in this study. B. Habitat Preferences Red swamp crayfish inhabit temporary lentic systems in its original home range, the lower Mississippi River Valley. Habitats are dry in the summer and crayfish survive in burrows. In the absence of predacious fish or in the presence of sufficient cover to protect the crayfish from predation, it establishes sustaining populations in both permanent lentic and lotic environments. As a result of introductions, its range now includes both the east and west coasts of the United States, Hawaii, Japan, Africa, Central America, and Europe (Huner and Romaire, 1978). Konikoff (1977) found that the main factors regulating natural P. clarkii populations were the hydrologic cycle and macrophyte densities and that predation can compound the effects of these factors in already unfavorable environments. Penn (1956) observed that in Louisiana most individuals of P. clarkii were found in shallow water (less than 15 inches deep) which was either clear or muddy, permanent, static, and exposed to full sunlight. Most collections were made in habitats with mud bottoms and aquatic vegetation. The aquatic plants most commonly associated with crayfish included bulrush (Scirpus sp.), cattails (Typha sp.), smarrweed (Pofygonum sp.), rush (Juncus sp.), and spiny naiad (Najas marina). Baker (1980) found that Procambarus was most often found in slow-moving water that was shaded, turbid, and had an abundance of aquatic vegetation for cover. Crayfish are most active at night (Penn, 1943). However, observation of the diurnal distribution of the red swamp crayfish in a clear stream in California showed a differential use of the center (exposed) and the bank (sheltered) portions of the stream by juveniles and adult crayfish. Juveniles were found more frequently in the open, unsheltered parts of the stream, and adults occupied the most sheltered areas (Beingesser, 1985). Witzig et al. (1983) found that P. clarkii densities were highest in shallow water and dense vegetation. When vegetation had decomposed and was not available as cover, crayfish showed no preference for deep or shallow areas. The authors suggested that terrestrial vegetation decomposed more slowly than aquatic vegetation and, therefore, served as a better source of cover and nutrients. C. Life History of the Red Swamp Crayfish Many details of the red swamp crayfish life history have been studied. The following is a summary of information on red swamp crayfish life histories in Louisiana presented by Penn (1956). In the winter, red swamp crayfish activity is restricted and most of the adults of both sexes over-winter in burrows in the drying marsh and swamp floors. Juveniles pass the winter in the water in or near the same localities, growing slightly. In January, with the advent of the pre-spring rains, the burrowing adults enter the water; shortly thereafter, the form U males change to form I (sexually active). In the spring, with increasing temperature and plant food supply, the juveniles grow rapidly, most of them reaching sexual maturity by May. 'Some copulation occurs in late spring but apparently many of the mature females retain sperm stored in their annul! from the previous summer and early fall. The first eggs are laid and carried by the females in late June and early July. Succeeding waves of egg-laying continue through August and early September. Most, if not all, of the year old males are in form I during the summer. Copulation occurs frequently in July and continues through September. Females which will live throughout the coming winter molt after their young have become free-swimming. The year's crop of juveniles appears in increasing numbers in late July and August. The mature population disappears at this time and apparently many of the females burrow. In September, most of the form I males and a few of the mature females leave the marsh by the tens of thousands and wander overland. Most of these crayfish fall prey to other animals; the remainder probably die since they are in extremely poor physical condition. The remaining adult females and large form II males burrow before the winter drought and cold sets in. The juvenile population remains in the water over winter. Studies of crayfish in Atchafalaya Basin, Louisiana, which contains water year round, resulted in the highest crayfish collection in May. By June, most crayfish were mature, and breeding occurred in the summer and fall. Youngof-the-year appeared in December, and their peak growth occurred in the spring (O'Brien, 1977). Red swamp crayfish have the potential, however, to reproduce during a large part of the year (Konikoff, 1977). This gives crayfish populations better tolerance of irregular variation in habitat conditions including variable hydrologic cycles and changing vegetation communities. Lowery and Mendes (1977) noted that although a portion of the red swamp crayfish population in an African lake was found to breed throughout the year, peaks of recruitment were found associated with rising water levels. Measurement of crayfish production in ponds with differing hydrological regimes (one with permanent water, one with irregular flooding events, and a pond with controlled flooding) showed that production was 32 times higher with a controlled flooding regime than at the other two sites (Faille, 1980). Changes in sex and sexual state (mature or reproductive to nonreproductive) ratios of crayfish populations occur seasonally and in relation to different nitrogen levels in ponds (Rhodes and Avault, 1987a). Huner (1983) found that development and molting of male crayfish occurred during different times of the year dependent on different flooding cycles. However, crayfish in all flooding cycles showed maturation peaks in the spring. Rhodes and Avault (1987b) found that peak egg production occurs during October of the female's first year. Only 17 percent of the females observed produced a second clutch of eggs the following April. Peaks in production observed around December are probably the result of second population of late maturing crayfish. D. Feeding Habits Penn (1956) reported that P. clarkii was observed feeding on the aquatic vegetation including Achyranthes, Jussiacea, Pofygonum, Sagittaria, Najas, Cfiara, Elodea, Potomogeton, and algae but not on Myriophyllum and Marsilea. Crayfish will also feed on periphyton, detritus, gastropods, and immature benthic insects (Lorman and Magnuson, 1978). E. Predation on Crayfish Saiki and Ziebell (1976) studied the effect of recently established crayfish populations on growth of stunted bass in an impoundment before the introduction of shad as additional forage. condition, Crayfish (Orconectes causeyi) densities," bass and food habits were evaluated over a one-year period. Young-of-the-year crayfish appeared in the spring, and maximum adult densities were measured in June. Bass 7.0 cm to 14.9 cm fed on crayfish in the spring, and larger bass consumed crayfish all year. Growth of older bass improved once crayfish were available as forage, and condition factors for the fish also increased. Stocking of crayfish is recommended to increase the forage base for bass in new impoundments. Saiki and Tash (1979) observed crayfish behavior in relation to predation by bass in laboratory and field situations. Predation rates on crayfish were examined in relation to density of plants as cover in plastic pools. Increased plant density significantly decreased predation rates. Size selectivity by bass for smaller crayfish also diminished at higher plant densities. Huner (1983) noted that experimental ponds containing green sunfish and crayfish (Procambarus spp.) showed significantly lower crayfish production levels than ponds containing only crayfish. II. DESCRIPTION OF THE STUDY AREA Lake Mead is located on the lower Colorado River where it forms the border between Clark County, Nevada, and Mohave County, Arizona (Figure 1). The regional climate is arid with an annual mean precipitation of less than 11.6 cm. The mean daily maximum temperature for July is 40°C and the mean daily minimum temperature for January is -1°C. Lake Mead was formed with the closure of Hoover Dam in 1935. The lake first reached maximum operating level in 1944, has a surface area of 660 km2 and a volume of 36 km3, and is one of the largest reservoirs in the United States. At maximum operating level, the shoreline of Lake Mead is 885 km, with a maximum length of 183 km and a maximum width of 28 km. It has a maximum depth of 180 meters and a mean depth of 55 meters. The all-time maximum operating level was reached during July of 1983. Since that time, the operating level has been successfully maintained at approximately 368 meters with annual variation of approximately four meters. In 1988, however, the lake level dropped approximately four meters between March and September, and remained at this low level for the rest of the year (Figure 2). Since the closure of Glen Canyon Dam in 1963 to form Lake Powell approximately 460 km upstream, most of Lake Mead has LIBRARY AUG 2 ! 1997 Buieau ol Reclamation Reclamation Service Center Lake Mead Arizona - Nevada Shoreline Elevation: 3 6 6 in lilue Point bay Roqei's Bay 00 W a t e r Barge Cove Fish Hatchery Cove Hoover Dam Figure 1. Map of Lake Mead. Grand Wash UJ __i LU Ld 1200^ 1195 J F M A M J J A S O N D MONTHS Figure 2. Lake Mead water elevations (in feet) fron January 1984 through December 1988. become extremely oligotrophic, with measured orthophosphorus levels as low as 0.001 mg/1 (Paulson and Baker, 1984). The dissected nature of the desert terrain flooded by Lake Mead has resulted in a cove and point topography. Many of the coves are located at the mouths of dry washes and receive inflows of runoff water containing nutrients and particulate organic matter during the infrequent rains. III. METHODS A. Literature Review A computerized abstract search was done to gather information on the red swamp crayfish. Approximately 200 articles were collected from numerous scientific publications and reviewed, and those considered important to the research were included in an annotated bibliography (Appendix A). B. Lake-wide Crayfish Population Sampling/Effects of Overton Arm Fertilization on Crayfish Populations A study to evaluate the population densities and distribution of red swamp crayfish in Lake Mead was started in the fall of 1986 (Peck et al., 1987). It was concluded from the lake-wide sampling (Figure 3) that red swamp crayfish were present throughout the lake and also occurred in some drainages entering the lake (Figure 4). Besides being widely distributed throughout the system, the population was found to be reproductively active. However, trap data indicated that the population density was extremely low. A catch per trap day (CPTD) value of one or less was common for the lake during the sampling period. Therefore, it is extremely difficult to study crayfish reproductive cycles, growth patterns, and 10 Muddy Arm Virgin Bowl Lake Meacl Arizona - Nevada Shoreline Elevation: 3(.Hi in Las Vegas Wash Hoover Dam Figure 3. Locations of red swamp crayfish sampling sites monitored between October 1986 and July 1987 Overton Wildlife Management Area Virgin River Procambarus clarkii LEGEND Shoreline Elevation 366 0 5 km in Lake Mead Rogers Spring Bighorn I s l a n d Las Vegas Bay Hemenway Harbor f«cl Cnvn» ?igure 4. Distribution of red swamp crayfish in Lake Mead from trap catches (October 1986-July 1987). habitat preferences by trapping or observation of naturally existing populations. Consequently, the trapping effort was reduced significantly from that outlined in the project proposal. An extensive trapping effort did, however, take place in the Overton Arm in October 1988 in order to investigate the effects of the Overton Fertilization Project on crayfish populations. In May and June 20,000 gallons of ammonium polyphosphate were put in the Overton Arm with the use of 200 volunteer boats. Traps were placed in the following Overton Arm areas: Mud Wash, Bark Bay, Fire Bay, Salt Bay, Glory Hole, Lime Wash, Calico Bay, and Roger's Bay. In each area a line comprised of 8 traps was set along the shoreline in 1-3 meter water depth in the following habitats: mud, brush, rocks/boulders, and spiny naiad (Najas marina). Results from this effort were compared to results gathered by previous investigators before the May 1988 Overton Arm Fertilization. C. Population Sampling and Life History Evaluation in Flamingo Wash Crayfish were trapped in Flamingo Wash four times in 1988: January, March, July, and November. Minnow traps with 4 cm openings were baited with chicken pieces and set at various locations in the wash. The traps were retrieved after 24 hours. Crayfish were sexed and carapace length was measured. Habitat conditions for each trapped area were recorded, including the presence of aquatic vegetation or other forms of cover. Trapping continued over several days until approximately 1,600 individuals were caught. These individuals were then used in the stocking program. 13 D. Lake Mead Stocking Program 1. Marking and Release Individuals trapped in Las Vegas Wash were placed in ice chests and transported to the laboratory where they were marked. Three different marks were used on each third of the trapped population. In January 1988, marks were placed on the left-most uropod and consisted of one hole for Lot 1, two holes for Lot 2, and a diagonal cut for Lot 3. Different uropods were marked with this code on each of the subsequent releases; in March, the second uropod from the left was used, in July, the third from the left was marked, and in November 1988, the right uropod was marked. Each marked group had an equal proportion of adults to juveniles and males to females. Crayfish were transported to a cove on Saddle Island in Boulder Basin of Lake Mead (Figure 5). This cove was selected because it had a wide range of habitats available for crayfish, including brush, mud, gravel, rocks, emergent plants, inundated terrestrial plants, and aquatic plants. In January, crayfish in Lot 1 were placed on a muddy bank with inundated terrestrial shrubs within the cove. Individuals in Lot 2 were placed near a small cattail stand, and individuals in Lot 3 were released on a gravel and rock embankment. As water levels receded during the year, the habitat in Lot 1 and Lot 2 release sites changed in subsequent introductions. In April, Lot 1 was released into a combination of inundated terrestrial and aquatic vegetation, with aquatic vegetation becoming the principal habitat in the last two introductions. Habitat at Lot 2's release area remained the same until the November release, 14 when aquatic Boulder Basin Figure 5. Location of study cove. 15 vegetation replaced the now desiccated cattail stand. The habitat at the Lot 3 release site remained the same for all introductions. 2. Monitoring Two days following the transplant, a series of ten 80 m long trap lines were established in the study cove (Figure 6). Eight standard size crawfish traps (41.9 cm long, 22.7 cm diameter, and a 5.7 cm diameter entrance) were used per line, and the traps were placed 10 m apart. Chicken pieces were used as bait. After the traps were set, a diving survey of the cove was made to map the general morphometry of the cove and the exact placement of each of the traps within various habitat components (plants, brush, mud, and rocks). Appendix B describes which trap lines/traps sampled a given habitat during the four monitoring periods. From this information, a' catch per unit effort (CPUE) was calculated for each habitat type represented in the study area. CPUE is calculated by taking the total number of crayfish caught per habitat type during a trapping period divided by the total number of traps used in that habitat during the same period. In addition, the crayfish movement patterns were determined by dividing the study cove into sections according to trap distances away from initial release sites. The distance categories used were: 0-40 m, 40-80 m, 80-120 m, 120-200 m, and 200+ m. The distances moved by crayfish from each lot's release site were figured for each monitoring date. Temperature, dissolved oxygen, conductivity, and pH were measured during each of the monitoring periods. 16 Figure 6. Trapline placeman! in- the study cove. The trap lines were retrieved 24 hours after their placement and the trapped crayfish were aged as adult or juvenile (carapace length less than 30 mm), sexed, and identified according to the trap they were in and their lot number. The traps were then re-established in their permanent sites, and crayfish were returned to the general area the trap line covered. Traps were set and retrieved every 24 hours for four days and then periodically thereafter or until returns reached 10 percent of the first day's catch. E. Predation by Game Fish NDOW conducted an annual creel census of the fish caught by anglers in Lake Mead from August 1987-August 1988. From this census, the stomach contents of each creeled game fish (largemouth bass and striped bass ) were analyzed and the percentage containing crayfish was determined. F. Habitat Improvement by Vegetation Introduction As part of the Cover Enhancement Project funded by NDOW, 1,200 sago pondweed tubers (Potamogeton pectinatus), an aquatic plant species, were placed in Roger's Bay in the upper basin of Lake Mead and 1,200 tubers were placed in Fish Hatchery Cove in the lower basin (Figure 1). Tubers were wrapped in cheesecloth bags and weighted with gravel. These bags were evenly distributed throughout the coves from the side of a boat to depths of between 2 and 15 meters. These introduction sites were monitored periodically for plant growth, percent cover, and fish use. Terrestrial and emergent vegetation species were also planted in the spring of 1988. 154 rooted woody plants and 344 direct woody cuttings were introduced at Bluepoint Cove in the upper basin in April and May 1988 (Figure 1). 241 rooted woody plants, 181 woody cuttings, and approximately 242 emergent plants were 18 introduced at Fish Hatchery Cove in April and May. Approximately 242 emergent plants were introduced at Roger's Bay in May 1988. In total, approximately 1,400 terrestrial and emergent plants were introduced in April and May. Species planted include: seepwillow baccharis (Baccharis glutinosa), willows (Salix gooddingii and 5. exigua), narrowleaf cattail (Typha angustifolia), and bulrush (Scirpus robustus). Sites were monitored bimonthly for survival and establishment of introduced plants. IV. RESULTS AND DISCUSSION A. Literature Review Appendix A contains an annotated bibliography of crayfish life histories, habitat requirements, effects and types of predation, and general crayfish ecology. B. Lake-wide Crayfish Population Sampling Trapping after the Overton Arm Fertilization Project resulted in a zero CPTD; the CPTD for the same area during the 1986 trapping effort was 0.05. Based on these results, it does not appear that the fertilization project had an effect on crayfish population densities in the Overton Arm. Therefore, it is concluded that the fertilization project did not have an effect on the benthic invertebrates or crayfish densities. C. Population Sampling and Life History Evaluation in Flamingo Wash Areas of Flamingo Wash, a natural drainage of the Las Vegas Valley, were sampled four times during 1988. Life history and population dynamics of the red swamp crayfish were determined. From these data, the highest crayfish densities were found in areas with a mud or silt bottom, some rocks, and extensive growth of cattails. Water depths ranged from 15 cm to 2 m, and there was a constant source 19 of slow moving water. Adult females represented approximately 44 percent of the total numbers caught for all the sampling dates except November-December when there was a 10 percent increase in the proportion of adult females (Figure 7). Adult females represented the dominant sex/age class for the 1988 sampling. In the April-May and July-August sampling periods, adult form I males represented 45 percent of the trapped crayfish, with 25 percent and 37 percent in the January-February and November-December periods. Adult form II (sexually inactive) males represented less than 10 percent of the trapped crayfish for all four sampling periods. In the January-February period, approximately 12 percent of trapped crayfish were represented by juvenile males and females, with their percentages decreasing to 1 percent in the November-December sampling period. The ratio of juvenile males to females was 1:1 during the sampling periods. It appears that there were no significant seasonal variations in the percentages of adult form II males and adult females; only the adult form I males and juveniles exhibited seasonal variations. Typically, the frequency of form II males increases in the winter when reproduction ceases and sexually active males (form I) molt into form IPs (Penn 1956). However, there was a decrease in the occurrence of form II males during the winter in Flamingo Wash. This inconsistency is either the result of reduced trapability of form IPs due to their susceptibility to predation, or a die-off of form Ps from low reproduction. The juveniles caught in January-February represent the release of over-wintered juveniles. These crayfish then matured and became adults by late spring/early summer. An additional increase in the number of juveniles should have occurred in July-August following the peak egg laying in June; this increase was not picked up in our sampling because of their reduced trapability in the summer due to increased predation. In conclusion, it appears that the life history of red swamp 20 100-, ESS Adult Female CZ1 Adult Male 1 tZ) Adult Male 2 CD Juvenile Male •I Juvenile Female 1 90 -j 8070- -£ 600) o 50i-_ 0) CL 40- 3020- s ^ _^ r^ \ ^ s \ \ 10- X n X R[ 1 Jan - Feb ^ X X X X ^ X X X X X ^ X X X X Rn. April - May \ ^ X Y s \ X X \ X X FL_ July — Aug X FL_ Nov - Dec Rgure 7. Percent of red swamp crayfish trapped in each sex/age class in Flamingo Wash during four trapping periods. 21 crayfish in Flamingo Wash is similar to the life history patterns reported in the literature. D. Lake Mead Stocking Program During each introduction, 1,600 marked crayfish were released into three areas of the study cove. An estimate of the native population in the study cove, life history trends of the crayfish in Lake Mead, habitat preferences, and movement patterns were determined from the subsequent monitoring. A total of 145 crayfish were recaptured for the January-February release, 661 for April-May release, 376 for July-August, and 575 for November-December releases (Table 1). Using Peterson's mark-recapture equation, Marked sample _ M sample + U sample M total M total + U total the native population of the study cove was estimated to be 65 individuals. The estimated number of native crayfish corresponds to a CPTD of 0.13 which compares to the outer Las Vegas Bay CPTD value of 0.10 calculated during the 1987 lake-wide sampling (Peck etal, 1987). In addition to low recapture numbers for each release, there was also a low percentage of hold-overs from previous releases. Only 6.2 percent of the crayfish stocked during the January-February release (Introduction 1) were recaptured during the April-May monitoring, with none being caught in the July-August or November-December periods. The April-May release had 2.9 percent held over into July-August, and only 0.3 percent of the July-August crayfish were recaptured during the November-December monitoring (Table 2). Low returns from previous releases are probably due to movement of crayfish out of the trap range. Also, it is possible that some of the crayfish died due to stress stemming from the introductions, predation, or natural causes. 22 Table 1: Number of red swamp crayfish recaptured after each release. Trapping continued until success was 10 percent of initial catch. Trapping Dates January-February April-May 145 July-August 661 376 November-December 575 Table 2: Percent recapture from previous releases. Trap Dates Jan-Feb April-May July-Aug April-May 6.2% N/A N/A July-Aug 0 2.9% N/A Nov-Dec 0 0 0.3% 23 1. Life History Percentages of adult females and form I males in the study cove were slightly different from those in the Flamingo Wash; however, they followed the same patterns over time (Figure 8). There were a lower precentage of form II males in April-May and July-August than the other stocking periods. period. The peak juvenile occurrence was in the July-August The life history trends exhibited by the red swamp crayfish in Flamingo Wash appear to parallel the trends exhibited in the study cove. 2. Habitat Preferences The most preferred habitat was emergent vegetation represented by a small stand of cattails (Typha angustifolia). A CPUE value of 20.0 was calculated for January-February (Introduction 1) release and 62.0 and 25.0 for the April-May (Introduction 2) and July-August (Introduction 3) releases (Table 3). By the fourth introduction (November-December), the cattail stand was desiccated, due to receding lake levels, and not available to the crayfish. The next most preferred habitat was inundated terrestrial shrubs represented by a small stand of seepwillow baccharis (Baccharis glutinosd) in another area of the cove. This habitat was only available during the first two introductions, with CPUE values of 2.5 and 14.0 respectively. Submerged aquatic vegetation was the third major habitat selected by red swamp crayfish, with sago pondweed (Potamogeton pectinatus) occurring in the April-May and July-August releases and spiny naiad (Najas marina) in the November-December release. Their calculated CPUE values were 12.75 and 6.83 for sago pondweed and 18.43 for spiny naiad. 24 100-| " 90 "I QQ j \ 70-£ ES Adult Female Cn Adult Male 1 CZ] Adult Male 2 i Juvenile Male •i Juvenile Female 60- (U \ (U S \s o 50- a. 4030- — n. s9 x X \ ^ \ S ^ Rn. Jan — Feb I ^ X X s X s s X s x n —— April — May X X X X X X X X $ I s x ^N X x s x 2010- 5 * X nfll July — Aug x Y PL. ' Nov — Dec Rgure 8. Percent of red swamp crayfish trapped in each sex/age class in the study cove during four trapping periods. 25 Table 3: Catch per unit effort of red swamp crayfish trapped in various habitats. POPE = sago pondweed (Potamogeton pectinatus} NAMA = spiny naiad (Najas marina). Habitat Jan-Feb Cattails 20.00 62.00 Inundated Terrestrial Shrubs 02.50 14.00 Submerged Aquatic Vegetation April-May July-Aug Nov-Dec 25.00 12.75 (POPE) 06.83 (POPE) 18.43 (NAMA) 05.58 Rocks 01.82 06.98 04.40 Brush 01.00 03.50 03.00 Gravel/Sand 00.82 03.41 01.64 01.29 Mud 00.41 02.07 01.27 01.22 26 Of the remaining habitat types sampled, rocks had CPUE values for four introductions of 1.82, 6.98, 4.40, and 5.58, respectively. Brush was available in the first three introductions with CPUE's of 1.0, 3.5, and 3.0. CPUE's in gravel/sand were 0.82, 3.41, 1.64, and 1.29. Mud was the least preferred habitat with a CPUE of 0.41 for January-February, 2.07 for April-May, and 1.27 and 1.22 for the last two introductions, respectively. The forms of vegetation selected by red swamp crayfish as preferred habitats offer a source of cover and food to the crayfish. By living at the bases of cattails and inundated terrestrials and within the dense mats formed by aquatic vegetation, the crayfish are better able to avoid predation by fish. In addition, the plant material provides an accessible food source for the resident crayfish. When vegetation is not available, rocks and brush are the next preferred habitats. However, they only provide a limited form of cover, and they also provide substrate for periphyton as a food source during certain seasons. Gravel/sand combinations and mud were the least preferred habitat because they offer virtually no cover or food sources to the crayfish. 3. Movement Patterns In addition to life history and habitat preferences of the stocked crayfish, the general movement patterns were also determined from the monitoring. During each of the introductions, there appeared to be a steady movement of crayfish out of the cove and into the surrounding areas. Figure 9 shows movements of crayfish released from the lot 1, 2, and 3 release sites for the January-February introduction. Crayfish released at the lot 3 location exhibited the most movement throughout the monitoring 27 CD ?J Number of crayfish trapped Nrrber of crayfish trapped I ro 01 ro V 0 I- o r J 1 \ Q 1 _ 00 8 ^ o o o T 8 8 8 b I o ' o 1 4 o 4 8 S 8 Number of crayfish trapped 8 5 1 NuT6sr of crayfish trapped 03 03 1 o CO o 03 03 o ro 8 \ \ 50\ \ i iS o 8 1 } I I ! 1 1 8 I a I 1 i i i | i 1 i • i NLmber of crayfish frcpped 5... .8 Number of crayfish trapped 03 03 CO ro ro 03 figure y. Number or red swamp craynsn trapped at various distances trom the Lot 1, Lot 2, and Lot 3 release sites during the first stocking period (January 26, 1989). (Continued) Lot 2 01-28-88 01-29-88 70 70 60 50 I 50 50 .-6 1 f. *° 8 •a *> 4O JO | 20 20 10 0 0-40 4O-80 80-120 120-200 20O+ 0-4O 4O-aD 02-02-88 8O-120 12O-200 20O+ 02-03-88 70 r /u tSSSSSs! htro 1 60- SO lillO 1 $ £ 40 1 % •*. 1 1 20 10 20 10 _, 0 0-40 4O-60 80-120 120-200 200+ 1 1 0-40 4O-80 02-12-88 ^ „ 1 ^ 1 80-120 120-200 1 200+ 02-25-88 /U 70 cssssss ESSSSS9 kilro 1 tx) 60 r "" ' - ' "" • ' ' • ' ' • " •"" • tttro 1 50 ^ 40 •^ 40 t g •5 -30 T 20 O ^ 0 , i 0-4O i 4O-80 i r 10 _, 80-120 120-200 i 0 1 1 1 1 200+ Ostcnce moved from htroobcticn site (meters) Distance movea from htrodjction site (meters) 29 1 O OJ \ 8 S S 8 8 Number of crayfish trcpped CO CO ro 2-3- 3 i; 2 f CD 3 O ^ i\ r \ \ I 1 1 O B | '-1 S S 8 Number of croyfish trcpped 8 Number of crayfish trapped co CO O CJ rn CO 0 (O 0 (0 V_ 1) I ?. I i 0Af 1a 8 8 S S Number of crayfish trapped ] • j i Number of croyfish trcpped . CD CD (O 0 i-« 1 8 03 co ro 1 ' I j = i 8 period, with no crayfish from this group being trapped on the last date. The majority of these crayfish were trapped between 80 and 200+ m from the release site. Crayfish released at the lot 2 location appeared to have the least amount of movement, with the majority of crayfish trapped in the 0-40 m range. Movement from the lot 1 location also appeared to be minimal. Lot 1 and 2 crayfish were released into areas with terrestrial or emergent vegetation, which may explain the slower movement out of these areas. Crayfish from lot 3 were released on a gravel and rock embankment which provided little food or cover to the crayfish. This presumably explains the greater amount of movement from this area. During the April-May release, lot 3 crayfish exhibited the same movement trends as in the first release, with the majority of crayfish caught between 80 and 200+ m (Figure 10). Crayfish released at the lot 2 location were mainly caught in the 0-40 m range except on the last date when they were found in the 80-120 m and 200+ m areas. Crayfish from lot 2 appeared to have the same movement trends in the April-May release as in the January-February release. Lot 1 crayfish showed greater movement in the April-May release than in the January-February release, with crayfish being caught in the 80-120 m and 200+ m areas on the last date. This is probably due to receding water levels which reduced the amount of inundated terrestrial vegetation available to the crayfish during the second introduction. In addition, some crayfish released during the first introduction in lot 1 and 3's locations were re-caught in all of the distance ranges during the April-May monitoring. However, first introduction lot 2 crayfish were found only in 0-80 meters from the release site. 31 OJ NJ o 5 8 iS S B Number of crayfish trapped Number of crayfish trcpped 8 03 T 03 I 3 6 8 03 CB 6 « 03 02 03 8 • 8 o 8 o en 5 Number of crayfish trapped a s a 8 a timber of crayfish o Ol I * Number of crayfish trapped 8 Number of crayfish trapped 10 OJ 5 S 8 8 8 S 8 Number of crayfish trcpped ti Number of crayfish trqpped 8 ro o 01 I CD TCD ro I 3 8 o i o i o o o Number of crayfish trapped o |!I Number of crayfish trapped CD ca O 02 03 CD 8 S 8 8 iS S S CD CD o m o en 1 II Number of crayfish trapped 8 Number of crayfish trapped figure J.O. iNumuej. OL leu swamp cj.dyj.ibJi Liappeu di_ Vdixous uxsLdiiceb iioin cut; J-.OL x, Lot 2, and Lot 3 release sites during the second stocking period (April 30, 1988). (Continued) Lot 3 05-02-88 05-03-88 70 60 50 20 10 0 0-4O 4O-80 80-120 120-200 20O+ 0-4O 4O-80 80-120 120-200 05-06-88 05-04-88 .-5 20O+ 70 70: 60 60! 50 50 4O X 40 g 30 20 20 10 10 0 0-4O 4O-BO SO-120 120-200 200+ 0-40 40-BO 80-120 120-200 200+ 05-26-88 05-12-88 70 70 60 60 50 * 40 40 ^o g -5 » 20 20 10 10 0 0-4O 4O-80 80-120 120-200 20O+ D'etnce moved from htrtxLictcn site I 0-4O T 4O-80 I 8O-120 120-200 200+ Distance movod fran niroduction site (meters) (meters) 34 Crayfish released in lot 3's location in July-August again exhibited the most movement, with the majority of the crayfish being caught in the 200+ m area (Figure 11). Lot 1 crayfish appeared to move out of the release site at a slower rate than in the previous two introductions; however, by the last monitoring date no crayfish from lot 1 were trapped. During July-August, a stand of sago pondweed was present in the lot 1 release site which may explain the slower outward movement, whereas the cattail stand in lot 2's site was gradually becoming desiccated due to receding water levels. This prompted more movement from lot 2 crayfish than had been exhibited in the first two releases. Second introduction crayfish were also caught during the July-August monitoring, with lot 1 and 2 crayfish caught from 0-120 meters away from their release sites, whereas lot 3 crayfish were found only in the 120-200 m range. Crayfish from lot 1 and 2 sites seem to stay in the release area longer than lot 3 crayfish which tended to move out of the study cove. Figure 12 shows movements of crayfish released at lots 1, 2, and 3 for the November-December introduction. Crayfish released at lot 1's site exhibited a minimal outward movement, with the majority remaining in the 0-40 m area. Lot 2 and 3 crayfish appeared to move steadily out of the release sites, with crayfish caught only in the 40-120 m and 200+ m areas for lot 2, and from 80-200+ meters for lot 3 on the last monitoring date. During the November-December release, lot 1's area had a dense bed of spiny naiad which offered ample cover for the crayfish, whereas lot 2 had a few scattered patches of spiny naiad and lot 3's area was comprised of gravel and rock. Apparently, neither area provided enough cover to hold the released crayfish in the area. However, lot 3 crayfish were the only ones 35 Figure 11. Number ot red swamp craynsn trapped at various distances rrom tne L,ot x, Lot 2, and Lot 3 release sites during the third stocking period (July 14, 1988). Lot 1 07-16-88 07-17-88 70 — mi^m 30 ' '" htTD 2 T) f '' | 50- ;5 40 -5 "S, 30 $ 3 20 i i 4O § D - htro 3 o A) 1 10 10| 0 i>-4O 4O-80 SO-120 120-200 1CXH- SSBI SB i 0-4O i -iO-80 SO-120 120-200 200+ D'stoxc moved trcm niroducticn site (meters) Oistance moved from ntrtxtictjon site Imetersl 07-18-88 07-19-88 70 70 60 hlro 3 60 50 I 40 ^-o ssar ggB r htro 3 5° ^ 40 8 30 •s 20 » I 20 10 10 i 0-4O i 4O-BO i 80-120 120-200 0-»O 200+ 40-flO 80-120 120-200 200+ Dstance moved from ntnxtictcn ste (meters) Distance mowed from ntrodicticn site (meters) 36 Figure 11. Number of red swamp crayrish trapped at various distances trom the Lot 1, Lot 2, and Lot 3 release sites during the third stocking period (July 15, 1988). (Continued) Lot 1 07-20-88 07-28-88 70 HBOI 00 60 50 -s 40 ! 1 DO •ft 40 1 30 1 i 20 30 20 10 10 0 u 0-40 4O-80 8O-120 120-200 litna 3 ^ 200+ i 0-4O 4O-SO 08-11-88 70 60 X 40 30 20 10 I 0--4O i l 80-120 120-200 l 200+ mcMsd hun ntrodjcticn stto (meters) Oistnce moved frcm ntrodjction site (meters) 0 ] i I 40-80 T T 80-120 120-200 200+ O'etnce moved fiun htnoduction ste (meters) 37 OJ 00 Number of crayfish trapped Number of crayfish Iropped CO 03 CD •s! o CB 03 OJ ?. o iff ;J ?i" 20- 6 6 8 8 8 S B Number of crayfish trapped IS of crayfish trcpped 8 o M CO co o ^J 5 C K H- f 0) H- 0 ft — (D in C (D & vO f <D CO 0 CO rt ^0 00 (D o 6 & S hS Number of crayfish tapped 8 Number of crayfish trapped CO CO O CD I 03 CO <s f M ?. o 8 £ 8 8 IS 6 8 Number of acyfish trapped 8 Number of crayfish trapped CD I I O 03 CO I1 (•0 O 1 8- S o a M CD CD in 0) • O O D rt HD c '•0 tt> tn *— ro d rt (t p- tn Ui • Co rt — ^o f oo o I-" - 3 1 .eo o o o — o — o OJ o £ o C£ o ro o t*i o ^ o ui Number of crayfish trapped c5 tO Number of crayfish trapped o co g ffl I o ^ tn O 8 . S »—* CD 03 CD o a IS S 8 Number of crayfish trapped 1 I CO CO CD CO ^ O 03 03 ffj O 05 f\ O Number of crayfish trapped CD O -• O e 3 rt H- O a c a 3 ft cu £0 C ft -"* O i f^ O ^£f C) I ! •fr O ^ O Nunnber of crayfish trapped Number of crayfish trapped S3 O "^ O Ui § T o o 6 8 6 8 8 is £ 8 Number of crayfish trapped 8 Nurrber of crayfish trapped i T CD CD 5r+1 ro o C fD 3 rt HD VI h1- o riyure xz. iMumoer o i rea swainp eraynsn trapped ac various distances rrom tne Lot 1, Lot 2, and Lot 3 release sites during the fourth stocking oeriod (November 15, 1982). Lot 1 11-17-88 11-19-88 70p TO J 40 ' b 'o | 20 10 0 4O-flO 80-120 120-200 200+ 0-40 4O-60 11-20-88 BO-120 120-200 200+ 11-29-88 70 r 70 60 60 40 I 30 ^ 10 10 0 0 0-4O 4O-80 80-120 12O-200 0-40 200+ 4O-80 Dctunce O'sbnce moved from htroductbn ste (meters) 12-13-88 70 B3SSB?;fq r>tro 4- 60 50 £ 40 ... o » ........ - 20 10 gBSu-— 0-40 4O-flO r^-^-^"" '• 80-120 120-200 200+ D'stcnce movea from ntroctiction ste (meters) 42 80-120 120-200 200+ frxum puxxkjctcn site i meters) O O T D O i o O Number of crayfish trapped O O CO !- o o Njrtier of crayfish trapped Number of crayfish trapped f*. o 03 03 CD I fO 03 03 I (0 O o o OJ o -fc o tn Number of crayfish trapped o y Number of crayfish trapped i i s 8 -a s 8 o — i o u> ^i o 03 03 co CD 03 3 o ft a (D BJ H 3 0 ngure i/. Numoer or rea swamp crayrisn trapped at various distances rrom tne Lot 1, Lot 2, and Lot 3 release sites during the fourth stocking period (November 15, 1982). (Continued) Lot 3 11-17-88 11-19-88 70 /o 60 60 BBH htro 3 50 .•6 - .-6 40i - - -._ hire 4 40 30 20j b-pr • i* 1 10 n iHi *i* i 10 0 0-4O 4O-80 i i 80-120 120-200 0 200+ 1 0-4O 4O-flO 11-20-88 itif'•'*• 20O+ 11-29-88 70 70 60 60 I 50 50 I 40 8 [ 80-120 120-200 «> X 20 101 0 4O-BO Dstoncc 80-120 120-200 200+ 0-40 fru 1 1 intrtxijctian site (meters) 40-flO 80-120 120-200 200+ O'stnoe mc*«d from ntrxxbcticn ste (meters) 12-13-88 70 60 50 40 30 20 10 0-40 4O-60 80-120 120-200 200+ Distance m&ad frcm ntroduction site (meters) 44 caught from the July-August release. They were found 120-200+ meters away from their release site. In conclusion, the crayfish in lot 3 showed more movement than in lots 1 and 2. This was probably due to an inadequate source of cover in the lot 3 release location. In addition, it is possible that the habitat and food sources in the study coves were inadequate to handle the number of crayfish released during each introduction, thereby contributing to the outward movement patterns exhibited by the released crayfish. From limnological data collected in the cove (Table 4), there were no signs of water quality problems (other than low productivity) which might have also caused the outward movement. E. Predation by Game Fish Wilde and Paulson (1989) found that in Lake Mead, the major component of striped bass diets during the spring of 1981 and 1982 were red swamp crayfish. However, the incidence of crayfish in game fish stomachs from August 1987 to August 1988 was extremely low. Of 975 striped bass stomachs analyzed, only 10 contained red swamp crayfish remains. Ninety-six largemouth bass stomachs were also analyzed for this period, and again only 10 contained crayfish remains. Even though recent stomach analysis indicates that red swamp crayfish are a minor component in the Lake Mead game fish diets, it is well documented in the literature that crayfish are utilized by bass as food. F. Evaluation of Plant Introductions Unpredicted low lake levels in 1988 resulted in the loss of many terrestrial plants. Seepwillow baccharis, however, had the highest survival rates. Aquatic plant tubers had 100 percent germination success and provided 70 percent cover for 45 Table 4: Mean water column temperature (C), dissolved oxygen (DO-mg-1), conductivity (EC-micro mho/cm) and pH in the study cove. Date/Time Temp. DO Cond. pH 1-28-88/1305 12.12 09.52 815 8.14 2-25-88/1215 12.23 10.52 816 8.21 5-12-88/1310 19.07 10.63 865 8.42 8-11-88/1250 27.98 10.21 867 8.54 46 fish by July 1988 (Haley et al., 1989). From the literature and the results of the introduction monitoring, it is apparent that vegetation plays a key role in habitat preferences of red swamp crayfish. Therefore, plant introduction into areas with crayfish would be beneficial to the population. V. CONCLUSIONS A. The current red swamp crayfish population density in Lake Mead is extremely low; however, they occur throughout the lake. B. The life history of red swamp crayfish in Lake Mead and in Las Vegas urban washes is similar to that reported in the literature. C. The population density of red swamp crayfish in Las Vegas urban washes is, in general, high. This is probably due to optimal environmental conditions including adequate habitat and food, a constant source of water, and no predation. D. Food preferences of the red swamp crayfish were reported by Penn in 1956. P. clarkii was observed feeding on the aquatic vegetation, including Achyranthes, Jussiacea, Potygonum, Sagittaria, Najas, Chara, Elodea, Potomogeton, and algae, but not on Myriophyllum and Marsilea. Crayfish will also feed on periphyton, detritus, gastropods, and immature benthic insects (Lorman and Magnuson, 1978). E. Based on the habitat preferences demonstrated by stocked crayfish, emergent vegetation, including cattails and aquatic vegetation (sago pondweed and spiny naiad), are important to red swamp crayfish on Lake Mead. F. This study indicates that stocked crayfish move away from a release site if preferred habitat is not available. Consequently, a vegetative cover enhancement project would provide habitat for stocked crayfish and help to maintain population levels in localized areas to facilitate breeding. G. Stocking crayfish in Lake Mead is relatively simple. However, the introduced populations do not become widely established and, therefore, do not provide sufficient forage to augment the shad population. 47 VI. LITERATURE CITED Allan, R.C. and B.C. Roden. 1978. Fish of Lake Mead and Lake Mojave. Base fisheries data, Boulder Canyon to Davis Dam. Nevada Dept. of Wildlife. 105 pp. Albert, E., and J.R. Baker. 1982. Food habits of sub-adult striped bass (Morone saxatilis) in Lake Mead, 1981-1982. Nevada Dept. of Wildlife. 13 pp. Baker, R.K. 1980. Distribution and abundance of three introduced crayfish species in the northern Sacramento Valley. M.A. Thesis. California State University. 40 pp. Beingesser, K.R., and N.H. Copp. 1985. Differential diurnal distribution of Procambarus clarkii (Girard) juveniles and adults and possible adaptive value of color differences between them (Decapoda, Astacidea). Crustaceana, 49(2)7:164-172. Haley, J.S., L.K. Croft, S.E. Leavitt, L.J. Paulson, and D.H. Baepler. 1989. Introduction and Enhancement of Vegetative Cover at Lake Mead. Lake Mead Limnological Research Center, submitted to Nevada Dept. of Wildlife. 85 pp. Huner, J.V. 1983. Crawfish, Procambarus spp., production from summer flooded experimental ponds used to culture prawns, Macrobrachium resenbergii, and/or channel catfish, Ictaluruspunctatus, in south Louisiana. Freshwater Crayfish 5:379-390. Huner, J.V., and R.P. Romaire. 1978. Size at maturity as a means of comparing populations of Procambarus clarkii (Girard) (Crustacea, Decapada) from different habitats. Freshwater Crayfish 4:53-64. Jonez, A., and R.C. Sumner. 1954. Lake Mead and Mohave investigations. Nevada Fish and Game Commission Final Report. D-J Project F-l-R, 186 pp, unpublished. Konikoff, M. 1977. Study of the life history and ecology of the red swamp crawfish, Procambarus clarkii, in the lower Atchafalaya Basin floodway. Final Report. U.S. Fish and Wildl. Serv., Univ. of Southwestern Louisiana. 81 pp. Lorman, J.G., and J.J. Magnuson. 1978. The role of crayfishes in aquatic ecosystems. Fisheries 3(6):8-10. Lowery, R.S., and A.J. Mendes. 1977. Procambarus clarkii in Lake Naivasha, Kenya, and its effects on established and potential fisheries. Aquaculture 11:111-121. O'Brien, T.P. 1977. Crawfishes of the Atchafalaya Basin, Louisiana, with emphasis on those species of commercial importance. Master's thesis. Louisiana State University. 77 pp. Paille, R.F. 1980. Production of three populations of red swamp crawfish, Procambarus clarkii, in southeast Louisiana. Unpublished. Master's thesis. Louisiana State University. 40 pp. Paulson, L.J., and J.R. Baker. 1983. The effects of limited food availability on the striped bass fishery in Lake Mead in Aquatic Resources Management of the Colorado River Ecosystem. pp. 551-561. Ann Arbor Scientific Pub. 48 Peck, S.K., W.L. Pratt, I.E. Pollard, L.J. Paulson, and D.H. Baepler. 1987. Benthic Invertebrates and Crayfish of Lake Mead. Lake Mead Limnological Research Center, submitted to Nevada Department of Wildlife. 84 pp. Penn, G.H. 1943. A study of the life history of the Louisiana red-crawfish, Cambarus clarkii Girard. Ecology 24(1): 1-18. Penn, G.H. 1956. The genus Procambarus in Louisiana (Decapoda, Astacidae). Rhodes, R.J., and J.W. Avault, Jr. 1987a. Changing sexual-state relationships of a population of red swamp crawfish Procambarus clarkii. NOAA Tech. Rep. NMFS. 0(47):51-52. Rhodes, R.J., and J.W. Avault, Jr. 1987b. A note on recruitment cycles for the red swamp crayfish Procambarus clarkii. NOAA Tech. Rep. NMFS. 0(47):53-54. Saiki, M.K., and J.C. Tash. 1979. Use of cover and dispersal by crayfish to reduce predation by largemouth bass. J. Ariz. Acad. Sci. 11:99-104. Saiki, M.K., and C.D. Ziebell. 1976. Some trophic relationships of the largemouth bass Micropterus salmoides (Laiepide) in a southwestern impoundment. J. Ariz. Acad. Sci. 11:99-104. Wilde, G.R., and L.J. Paulson. 1989. Food habits of subadult striped bass in Lake Mead, Arizona-Nevada. The Southwestern Naturalist. 34(1). Witzig, J.F., J.W. Avault, Jr., and J.V. Huner. 1983. Crawfish, Procambarus clarkii, growth and dispersal in a small south Louisiana pond planted with rice, Oryza sativa. Freshwater Crayfish 5:331-343. 49 Appendix A ANNOTATED BIBLIOGRAPHY TO CRAYFISH ECOLOGY Abrahamsson, S. 1973. Methods for restoration of crayfish waters in Europe--the development of an industry for production of young Pacifastacus leniusculus DANA pp.204-210 in Freshwater Crayfish (S. Abrahamsson, ed). Studentlitterautur. Lund, Sweden. Pacifastacus leniusculus was introduced to Sweden after a plague destroyed the native Astacus astacus. Over 1/2 million juvenile Pacifastacus leniusculus were raised in hatcheries and released into Swedish lakes. Juveniles stocked in early spring were reproducing within a year. After one year, 59 percent of juvenile introductions were successful as compared to 30 percent success rates with adult crayfish introductions. One year after a release of 1,000 juveniles in each of two ponds, recapture rates were 21 percent and 33 percent in spite of high numbers of fish predators. Abrahamsson recommended the quantity of introduced crayfish should not be too low because low densities will reduce the chance of mating. Introductions should be repeated for some years to ensure the optimum development of the crayfish populations. Adeboye, Duro. "The Crayfish Condition Factor: A Tool In Crayfish Research." Freshwater Crayfish Abstract 5. A "crayfish constant, K" and a "crayfish condition factor, F" were calculated for 147 specimens of Procambarus astacus astacus. It was found that the "crayfish constant" was constant for all sizes and sexes in different molting cycle stages. The "condition factor" was designed to facilitate comparative physiological studies of various crayfish species from different laboratories. The ratio of body weight to the products of linear measurements were used to calculate the "crayfish constant." K= Body Weight Total Length X Carapace Length X Carapace Width The unit for K is gm/cm3. The "condition factor" was derived from the product of K and the assigned numerical value of the moulting cycle stage. F= K X Moulting Stage The unit for F is gm stage/cm3. Avault, J. W. 1973. "Crayfish Farming in the United States." Freshwater Crayfish 1:240. There are currently three types of ponds being used for crayfish farming: rice field, wooded, and open ponds. The ponds are flooded, a practice which provides the crayfish with food and cover sources (inundated vegetation). Crayfish diets consist mainly of plant material, small organisms, and periphyton. Rice fields and open ponds appear to be the most effective, due to an ample food supply available to the crayfish and do not exhibit an oxygen depletion or low pH~unlike the wooded ponds. Crayfish farming has steadily been increasing because they are an excellent food source for game fish. A-2 Baker, R. K. 1980. Distribution and abundance of three introduced crayfish species in the northern Sacramento Valley. M.A. Thesis. California State University. 40 pp. The distribution, relative abundance, and habitat of the crayfish Pacifasticus leniusculus, Procambarus clarldi, and Orconectes virilis are presented as examined in areas associated with the Sacramento River in California. Several physical habitat characteristics were measured and correlated, including the nature of the water course, vegetation, cover, and substrate, as well as dissolved oxygen and temperature. Distinct patterns of habitat segregation were found between species. Due to the method of collecting crayfish (by hand from under rocks), information on P. clarldi is mainly presented for generally sub-optimal habitat types. Beingesser, K. R., and N. H. Copp. 1985. Differential diurnal distribution of Procambarus clarldi (Girard) juveniles and adults and possible adaptive value of color differences between them (Decapoda, Astacidea). Crustaceana, 49(2)7:164-172 Diurnal distribution of Procambarus clarldi was observed in a clear stream in California. Differential use of the center (exposed) and the bank (sheltered) portions of the stream by juveniles and adults was found. Juveniles (<60mm total length, mottled green and brown in color) were found more frequently in the open, unsheltered parts of the stream. Adults (darker coloration-red) occupied the most sheltered areas most frequently. The lighter color of juveniles may render them less susceptible to diurnal predators. Black, J.B., and J.V. Huner. Breeding your own crayfish stock. (No references-miscellaneous paper. This is a summary of Procambarus clarldi life cycle under laboratory conditions at 23-26°C. Chlorinated tapwater was used (hardness must be 50 ppm calcium at minimum). Crayfish were fed alligator weed (altemanthera), Elerdea or rotted, soaked hardwood leaves, and dried catfood. Males were in breeding form 8-9 months of the year. Females produced eggs 2-3 times a year. Young matured at about 7.5 cm in 6-8 weeks and were ready to breed within about 4 months. Brinck, P. 1977. Developing crayfish populations. Freshwater Crayfish 3:211-228. A summary of crayfish (Pacifastacus leniusculus) introductions conducted over seven years in Europe is presented. Rationale for stocking strategies and follow-up studies is included. The success of establishing populations increased as the stocking frequency increased. However, the author noted that determination of reproduction in the lake can be tricky when stocking has been repeated. Brown, D.J., and J.M. Brewis. 1978. A critical look at trapping as a method of sampling a population of Austropotambius pallipes (Lereboullet) in a mark and recapture study. Freshwater Crayfish 4:159-164. This study compares results from trapping to hand capture of crayfish. Population estimates were made by mark recapture using both collection methods. Although both methods exhibited some bias, trapping showed the greatest amount, underestimating population size three-fold. Some alternatives are discussed, including alternation of sampling methods when using mark A-3 recapture to estimate crayfish population size. Chien, Y., and J.W. Avault, Jr. 1983. Effects of flooding dates and type of disposal of rice straw on the initial survival and growth of caged juvenile crayfish, Procambarus clarkii, in ponds. Freshwater Crayfish 5:344-350. Caged crayfish were placed in ponds flooded in September or October with three different types of rice straw sewage. Crayfish were fed a pelleted ration, so that differences in growth were independent of food. Dissolved oxygen and temperature was monitored in all treatments. Higher DO and temperatures enhanced growth but resulted in lower survival. Growth of juvenile crayfish (14.5 mm carapace) averaged 8.7-9.3 mm over the 4-week study period. Chien, Y.H., and J.W. Avault, Jr. 1983. Effects of flooding dates and disposals of rice straw on crayfish, Procambarus clarkii (Girard), culture on rice fields. Aquaculture 31:339-359. Population dynamics and production of crayfish cultured in ponds in Louisiana was followed to assess the effect of early (September) and late (October) flooding dates in conjunction with three methods of rice straw (food source) disposals. Growth of periphyton in the ponds was monitored during the study, as well as physical conditions. The discussion of this study highlights the interrelationships of factors such as temperature, dissolved oxygen, structure (cover), and food type on survival, growth, and density of crayfish in a situation with relatively high densities. Clark, D.F., and J.W. Avault, Jr. 1975. Effects of feeding, fertilization, and vegetation on production of red swamp crayfish, Procambarus clarkii. Freshwater Crayfish 2:125-138. This study evaluates the growth and survival of juvenile crayfish stocked in metal pools during summer and winter. The effect of 2 feed types, plantings of smartweed (Potygonum) and alligator weed (altemanthera philoxeroides), addition of fertilizer, and control (no feed added) on production was examined. Survival of crayfish was similar in all treatments. Growth and production was highest in feed treatments: average weight crayfish 21-22g. Average growth and production occurred on both the plant species; and when fertilizer only was added to the pools, the average weight of the crayfish was 13 g. Crayfish apparently are able to survive on phytoplankton. Collins, N.C., H.H. Harvey, AJ. Tierney, and D.W. Dunham. 1983. Influence of predator density on trapability of crayfish in Ontario Lakes. Can. J. Fish. Aquat. Sci. 40:1820-1828. Density and trapability of crayfish populations were measured in several Ontario lakes. Trap results were compared to counts made by SCUBA divers to evaluate crayfish density. Density of predatious fish species was also measured for each lake. Trap results did not always accurately reflect crayfish densities. In some lakes with higher fish densities, crayfish densities were higher than reflected by trap returns. Therefore, high predator density not only decreased crayfish abundance in some lakes, but in other lakes affected trapability of crayfish. Laboratory observations of crayfish from lakes which differed in predator abundance confirmed that a behavioral change resulted in lower trap catches. Crayfish from a lake with high bass densities were less active than crayfish from a lake without bass. A-4 Dye, L., and P. Jones. 1975. The influence of density and invertebrate predation on the survival of young-of-the-year Orconectes virilis. pp. 529-538 in Freshwater Crayfish 2 (J. W. Avault, Jr., editor). Louisiana State University Division of Continuing Education. Baton Rouge, LA. The effects of density and invertebrate predation on juvenile Orconectes virilis was examined in two small Michigan lakes in 1973. Survival of young-of-the-year crayfish in field enclosures at densities of 10.8/m2, 43/m2, and 172/m2 with no predators was significantly higher in the 10.8/m2 enclosure. There was no significant difference in survival of crayfish in 43/m2 and 172/m2 densities. Three invertebrate predators, Aeshnids, Notonectids, and Aeshnids significantly reduced crayfish populations with 4-6 mm carapace lengths but not with 6-10 mm carapace lengths. Predation by adult crayfish and Notonectids was not significant on either size of crayfish. Fitzpatrick, J.F., Jr. 1977. The statistical relationships of different techniques of measurements in a crayfish species. Freshwater Crayfish 3:471-479 (O.V. Lindquist, editor). University of Kuopio, Finland. This study examined three methods of carapace measurement: standard carapace length, postorbital carapace length, and maximum carapace length. Standard carapace length and maximum carapace length were better for statistical analysis than postorbital carapace measurements. The ratios of each type of carapace length to areola length (an important taxonomic ratio) were determined and did not correlate as well as unsealed carapace length data. Thus, either standard carapace length or maximum carapace length is the best method for measuring crayfish length. Furst, M. 1977. Introduction of Pacifastacus leniusculus (Dana) into Sweden: Methods, results and management. Freshwater Crayfish 3:229-247. A summary of procedures and considerations made during the initial introduction of crayfish from the United States to Sweden. Both adult and juvenile crayfish were stocked in various numbers and follow-up trapping was done to evaluate success in different areas. Adults tended to move large distances after transplant (up to 600 M). If adult crayfish were held in cages for four days, they appeared to "imprint" on the area and showed less movement (maximum distance 25 M). However, stocking lakes with adults was stated to be the most effective and least expensive method. Hazlett, B.A., and D. Rittschof. 1985. Variation in rate of growth in the crayfish Orconectes virilis. J. Crust. Biol. 5(2):341-346. Growth of individual adult crayfish (Orconectes virilis) living in a Michigan stream was followed by mark-recapture. 1,021 individuals were followed for a summer. Thirty-six crayfish were measured one year later. Growth per molt was quite variable, ranging from 0-9 mm per molt. Crayfish molted as many as three times during the summer. The growth rate of crayfish in this stream is higher than other populations. Unlike other populations, crayfish in this study frequently burrow. It is hypothesized that the higher growth rate may result from the burrowing behavior, since this lowers the effective density as well as providing crayfish with roots as an additional food source. A-5 Huner, J.V. 1975. Observations on the life histories of recreationaily important crawfishes in temporary habitats. Crayfish (Procambarus clarkii and P. a. acutus) were sampled monthly with dip nets and traps for a full year. Sampling was done in shallow (.15-.30 m) drainage ditches in southern Louisiana, which contained Altemanthera philoxeroides, Pofygonum, and grasses. Juvenile (less than 30 mm total length) crayfish numbers peaked in August and January for P. clarkii. Form I (mature) males showed two peaks, in August and in April. This data illustrates that P. clarkii is capable of multiple generations in habitats where drought conditions do not prevent them from reproducing during early summer. Information is also presented on minimum time required for crayfish to mature, sizes at maturity, and success of a limited mark-recapture study. Huner, J.V., and J.W. Avault, Jr. 1976. Sequential pond flooding: A prospective management technique for extended production of bait-size crawfish. Trans Am Fish Soc. 105(5):637-642. Ponds in Louisiana were stocked in May, drained in June, and reflooded the first of September, October, or November. From the middle of November, crayfish were sampled by the use of dip nets and traps biweekly through May, measured, sexed, and released. Maturity of males was also recorded, molt stage was determined, and crayfish over 4.5 cm were marked. Life history differed between flooding dates. Early maturation (October) occurred in ponds flooded in September and October but not in November. All ponds exhibited maturation peaks in spring. Higher proportion of premolt (molt within two weeks) crayfish were caught in dip nets than in traps. Densities of non-juvenile crayfish were estimated at 9.4-11.7 per square meter. Regular harvesting of crayfish populations is recommended to enhance growth, especially when densities are high and food potentially limiting. Huner, J.V. 1977. Introductions of the Louisiana Red Swamp Crayfish, Procambarus clarkii (Girard); an Update. Freshwater Crayfish 3:193-202. Procambarus clarkii occurs naturally in the south-central United States and parts of northern Mexico. However, it has been successfully introduced into California, Nevada, Hawaii, Japan, parts of Africa, and Central America. This species is adapted to temporary, 'lentic environments and can produce two generations annually, which has helped its establishment in new areas. P. clarkii is proving to be a serious problem in some introduced areas, due to its damage of irrigation levies and young plant shoots. In areas where crayfish are harvested for commercial and fishery uses, the populations are kept in check, and a problem has not occurred. Huner, J.V., and R.P. Romaire. 1978. Size at maturity as a means of comparing populations of Procambarus clarkii (Girard) (Crustacea, Decapada) from different habitats. Freshwater Crayfish 4:53-64. Size frequency distributions of mature (Form I) male crayfish are presented for temporary (ditch and pond) and permanent (basin/lake) water habitats. Differences in factors such as food availability, water quality, predation levels, and harvesting frequency on the population dynamics of crayfish in these habitats are discussed. Size at maturity (ranging from 54-134 mm total length) was smallest for males in roadside ditches, intermediate in most ponds, and largest A-6 in optimal basin (lake/floodway) areas. Life history of Procambarus clarkii is reviewed in relation to data on size and variation in size of mature males due to various habitat characteristics. Huner, J.V. 1983. Crawfish, Procambarus spp., production from summer flooded experimental ponds used to culture prawns, Macrobrachium rosenbergii, and/or channel catfish, Ictalurus punctatus, in southern Louisiana. Freshwater Crayfish 5:379-390. In contrast to the traditional practice of draining crayfish ponds during the summer months, production of crayfish in ponds flooded during the summer was examined. In addition, the polyculture of crayfish with fish and prawns was examined. Crayfish did complete their lifecycle even though ponds were not drained during the summer. However, production of crayfish was lower. Control ponds (ponds with only crayfish) were not included, so effects of flooding dates and species interactions cannot be differentiated. However, ponds containing green sunfish (known predator of juvenile crayfish) showed significantly lower production levels. Jaspers, E.J.M. 1969. Environmental conditions in burrows and ponds of the red swamp crawfish, Procambarus clarkii (Girard), near Baton Rouge, Louisiana. Master's thesis. Louisiana State University. 47 pp. A comparison was made of a number of factors measured in crayfish irregular flooding events and a pond with regular, controlled flooding. Annual production was 32 times higher for the site with controlled flooding than for the other two sites. A number of environmental parameters were measured at each site and found to be similar between sites. Therefore, several aspects related to the hydrologic regime may account for the difference in production by crayfish. Correlation of crayfish densities to physical-chemical parameters is also included. The highest number of crayfish was sampled in May. By June, most of the crayfish were mature. Breeding occurred in summer and fall. Young-of-year appeared in December, and peak growth occurred in the spring. The single generation exhibited by Procambarus clarkii in this study appears to be related to a low-water period lasting from August to October. Konikoff, M. 1977. Study of the life history and ecology of the red swamp crawfish, Procambarus clarkii, in the lower Atchafalaya Basin floodway. Final Report. U.S. Fish and Wildl.- Serv., Univ. of Southwestern Louisiana. 81 pp. This is a 15 month study of crayfish from six different areas, including swamp and lake habitats in southern Louisiana. Description of Life .history includes relative abundance, timing of reproduction (from field collections and examination of gonads), and growth rates. Differences in habitats were noted and environmental conditions were monitored. Crayfish were sampled by several methods, including examination of burrows, night lighting, and a mark recapture study to document crayfish movements. Examination of stomach contents was also done. Although large differences occurred between habitats, crayfish generally reproduced in late summer to fall. However, results in this study illustrated this species has the potential to reproduce during a large part of the year. This variability and the non-synchrony of the life cycle allows them better tolerance of irregular variation in habitat conditions. The main factors noted regulating natural populations were the hydrological cycle and macrophyte densities. Predation levels can compound the effects of these factors in already unfavorable situations. A-7 Lowery, R.S., and A.J. Mendes. 1977. Procambarus clarkii in Lake Naivasha, Kenya, and its effects on established and potential fisheries. Aquaculture, 11:111-121. The reproduction and distribution of crayfish in an African lake was examined four years after introduction. The population reproduced successfully and its distribution expanded. Although a portion of the population was found to breed throughout the year, peaks of recruitment were found associated with rising water levels. Unfortunately, little information on other physical/chemical parameters is presented. However, females in berry were reported from open water as well as from burrows. Mark recapture was used to estimate densities and provided limited movement data. Observations show that plant populations have declined in areas where crayfish are abundant. Crayfish have become an important food base for black bass (Micropterus salmoides) in this lake, but may also negatively impact certain aspects of the fisheries operations. Mason, J.C. 1979. Effects of temperature, photoperiod, substrate, and shelter, growth, and biomass accumulation on juvenile 82, P.J. Laurent, editor. Institut National de la Recherche Agronomique, Thoron-les-Bains, France. The objectives of this study were 1) to evaluate the relative importance of temperature and photoperiod in determining growth and survival of stage two Pacifastacus leniusculus in intensive culture; and 2) to evaluate substrate, shelter, light, and diet on growth and survival rates in intensive culture. The best growth rates were seen with high temperature and short photoperiod, with temperature being the predominating influence. Survival was inversely related to growth, high temperature, and short photoperiod. Average growth rate, biomass, and survival were not affected by diet. Survival and biomass were significantly affected by substrate. In a comparison of high density (260 crayfish/m2) vs. low density (130 crayfish/m2) and substrate, mortality was density-dependent on bare floor but density-independent with pebbled floor. Burrow use was density-dependent and increased the probability of survival. Melancon, E., Jr., and J. Avault, Jr. 1977. Oxygen tolerance of juvenile Red Swamp Crayfish, Procambarus clarkii (Girard). Freshwater Crayfish 3:371-380. LC50 values for acute dissolved oxygen depletion were determined for two size classes of juvenile crayfish (total lengths of 9-12 mm and 31-35 mm). The LC50 value for 96 hours of newly released 9-12 mm juveniles ranged from 0.75-1.10 ppm oxygen. The LC50 value for 96 hours for 31-35 mm juveniles was 0.49 ppm oxygen. Crayfish in the 31-35 mm size class that were molting had higher mortality rates than intermolting juveniles. O'Brien, T.P. 1977. Crawfishes of the Atchafalaya Basin, Louisiana, with emphasis on those species of commercial importance. Master's thesis. Louisiana State University. 77 pp. This is a study of the life history of several species of crayfish, including Procambarus clarkii collected from a basin which contains water year-round and yields a large number of crayfish commercially. Correlation of crayfish densities to physical-chemical parameters is also included. The highest number of crayfish was sampled in May. By June, most of the crayfish were mature. Breeding occurred in summer and fall. Young-of-year appeared in December, and peak growth occurred in the spring. The single generation exhibited by Procambarus clarkii in this study appears to be related to a low-water period, lasting from August to October. A-8 Faille, R.F. 1980. Production of three populations of red swamp crawfish, Procambarus clarkii, in southeast Louisiana. Unpublished. Master's thesis. Louisiana State University. 40 pp. Production of crayfish populations was measured from three areas exhibiting different hydrological regimes: one with permanent water, an area with irregular flooding events, and a pond with several methods, including examination of burrows, nightlighting, and a mark-recapture study to document crayfish movements. Examination of stomach contents was also done. Although large differences in abundances occurred between habitats, crayfish generally reproduced in late summer to fall. However, results in this study illustrate that this species has the potential to reproduce during a large part of the year. This variability and the non-synchrony of the life cycle allows them better tolerance of irregular variation in habitat conditions. The main factors noted regulating natural populations were the hydrological cycle and macrophyte densities. Predation levels can compound the effects of these factors in already unfavorable situations. Paille, R.F. 1980. Production of three populations of red swamp crawfish, Procambarus clarkii, in southeast Louisiana. Unpublished. Master's thesis. Louisiana State University. 40 pp. Production of crayfish populations was measured from three areas exhibiting different hydrological regimes: one with permanent water, an area with irregular flooding events, and a pond with regular controlled flooding. Annual production was 32 times higher for the site with controlled flooding than for the other two sites. A number of environmental parameters were measured at each site and found to be similar between sites. Therefore, several aspects related to the hydrologic regime may account for the difference in production by crayfish. Differences between sites in food availability, predation rates, flushing rates, and the effect of irregular changes of water levels on reproduction itself were considered as possible factors. Of these, it was concluded that predation may be the most critical factor limiting crayfish population. Payne, J.F. 1978. Aspects of the life histories of selected species of North American crayfishes. Fisheries Vol. 3, No. 6, pg. 5-8. Female crayfish secrete a translucent mucilaginous sac where the eggs and sperm are mixed and where fertilization eventually occurs. The sac wall begins to harden and cements the eggs to pleopods. This occurs mainly during late summer and early fall in Procambrius clarkii. Egg development is dependent on temperature, photoperiod, water circulation, and hormonal balances within the female. It is not unusual if 40 percent of the eggs fail to hatch due to infertility, damage resulting in a fungal attack, and the failure of the eggs to attach to the pleopods. The hatched crayfish remain attached to the female's pleopods during the first and second instars, which lasts approximately 19 days. Third instar crayfish venture from the female for short periods of time. P. clarkii females produce a brooding phermone which allows a chemical communication to exist between the female and juveniles. Maturation rates are highly variable and are dependent on availability of food, water quality, and temperature. If conditions are right, P. clarkii juveniles may reach adult size within two to three months, and two generations may be produced within a single year. Penn, G.H. 1956. The genus Procambarus in Louisiana (Decapoda,Astacidae). A summary of life history and ecology based on data from taxonomic collections and early life history studies of Procambarus clarkii in Louisiana are presented. A single generation life cycle A-9 is described. Also included is a list of associated plant species and records of plants P. clarkii has been observed feeding on. Pollard, I.E., S.M. Melancon, and L.S. Bakey. 1983. Importance of bottomland hardwoods to crawfish and fish in the Henderson Lake area, Atchafalaya Basin, Louisiana. Wetlands, 3:1-25. Distribution and abundance of several fish species, young-of-year largemouth bass, crawfish, and zooplankton were in areas including and associated with flooded woodlands in Louisiana. In addition, several physical/chemical parameters were monitored; stomach contents of young-of-year largemouth bass were examined; and a mark-recapture study to estimate crawfish abundance was done. The highest concentrations of crawfish were found associated with the edge of floodwaters, probably resulting from increased availability of food. However, greater concentrations of crawfish burrows were found closer to permanent water. Young-of-year crawfish were observed to be important in the diets of adult largemouth bass. Price, J.O., and J.F. Payne. 1979. Multiple summer molts in adult Orconectes neglectus chaenodactylus Williams. Freshwater Crayfish 4:93-104. A mark-recapture study was used to determine molting cycle, growth rate, and the population size class composition of Orconectes neglectus chaenodactylus in Arkansas. Over 752 animals were marked on the carapace or abdomen using a heated dissection probe. The article illustrates the numbering system. The mean minimum size for adult males and females was 13.5 mm (carapace length). Females molted approximately two weeks after release of young and then 2 to 3 times during the summer. Males molted from Form I to Form II during April and May, then molted twice in the summer, remaining in Form II. The males then molted to Form I in late October and November. Adult crayfish did not molt during winter. Population size class composition of reproductive size crayfish was dominated by two-year-olds in the spring. As the average life span of this species is 2.5 years, by fall many of the adults died, leaving the sub-adult age/size class predominating the population. Rhodes, R.J., and J.W. Avault, Jr. 1987. Changing sexual-state relationships of a population of Red Swamp crawfish Procambarus clarkii. NOAA Tech Rep NMFS. 0(47): 51-52. Changes in sex and sexual state (mature or reproductive to non-reproductive) ratios of crayfish populations were found to occur seasonally and in relation to different nitrogen levels in ponds. External determination of mature (inseminated) females and mature-form males is described. The changes in these ratios are discussed in relation to crayfish population dynamics as a predictive tool for evaluating crayfish growth and reproductive responses to different conditions. Rhodes, R.J., and J.W. Avault, Jr. 1987. A note on recruitment cycles for the Red Swamp Crawfish Procambarus clarkii. NOAA Tech Rep NMFS. O(47):53-54. Oviposition by Procambarus clarkii was examined over an eight-month period in the laboratory for comparison with field observations of recruitment. Females which layed eggs were followed to determine their ability to produce a second clutch. Peak egg production occurred during October of the first year with 56 percent of the females releasing eggs. Only 17 percent of the A-10 females produced a second clutch of eggs the following April. G mortality and deformity due to molting was noted in females which had oviposited once. Production peaks in October and April agree with field recruitment. It was concluded that a third peak observed in field populations during December must result from a second population of crayfish which mature later. Romaire, R.P., and V.A. Pfizter. 1983. Effects of trap density and diel harvesting frequency on catch of crawfish. N. Am. J. of Fish. Manage. 3:419-424. The relationship of trap density and frequency of harvest to catch of crayfish was examined in commercial crayfish ponds. Crayfish catch increased with higher densities of traps. The optimal frequency of harvest was achieved with two six-hour (diurnal) sets and one twelve-hour (nocturnal) set. Although increased trap effort yielded greater numbers of crayfish in this study, when crayfish densities were high, this relationship may be dependent on densities of crayfish as well as other factors. Saiki, M.K., and C.D. Ziebell. 1976. Some trophic relationships of the largemouth bass Micropterus salmordes (Laiepide) in a southwestern impoundment. J. Ariz. Acad. Sci. 11:99-104. This study evaluated the effect of recently-established crayfish populations on growth of stunted bass in an impoundment before the introduction of shad as additional forage. Crayfish (Orconectes causeyi) densities, bass condition, and food habits were evaluated over a one-year period. The macrophyte Myriophyllum exalbescens grew abundantly along the shore and served as crayfish forage. Young-of-year crayfish appeared in the spring, and maximum adult densities were measured in June. Bass from 7.0 cm to 14.9 cm fed on crayfish in the spring, and larger bass consumed crayfish all year. Growth of older (stunted) bass improved once crayfish were available as forage, and condition factor of fish also increased by the end of the study. Since shad had not been introduced yet, it was concluded that crayfish alone could improve growth of largemouth bass. Stocking of crayfish is suggested to improve forage bases for bass in new impoundments. Saiki, M.K., and J.C. Tash. 1979. Use of cover and dispersal by crayfish to reduce predation by largemouth bass. J. Ariz. Acad. Sci. 11:99-104. Observation of crayfish behavior in relation to predation by bass was done in lab and field situations. Predation rate on crayfish was examined in relation to density of plants as cover in plastic pools. Increased plant densities significantly decreased predation rates. Size selectivity by bass for smaller crayfish also diminished at higher plant densities. Trapping of crayfish in a southwestern lake which contained water milfoil [primary cover for crayfish] and largemouth bass was conducted to examine seasonal distribution of crayfish. Greater numbers of crayfish were found in vegetated areas. However, crayfish dispersed throughout the lake when plants died back. Therefore, multiple behavioral interactions may be responsible for achieving balanced population levels with this particular food web. Sheppard, M.F. 1974. Growth patterns, sex ratio, and relative abundance of crayfishes in Alligator Bayou, Louisiana. Master's thesis. Louisiana State University. 54 pp. Life histories of six species of crayfish, including Procambarus clarkii, were studied in a swampy A-ll area subjected to periodic flooding in southern Louisiana. Crayfish were sampled by dip netting and seining monthly over a two-year period. Size, frequency, distribution, and resulting growth patterns of 2,068 P. clarkii sampled during the study are presented. There appears to be one generation per year, young first appearing as early as September, slow growth occurring during the winter, increasing in April to June until the majority of the population was adult size. It was also observed that Procambarus spp. readily moved into the inundated floodplains during high water, allowing access to additional food sources. Somers, K.M., and D.P.M. Stechey. 1986. Variable trapability of crayfish associated with bait type, water temperature, and lunar phase. Am. Midi. Nat. 116(l):36-44. Comparison of trap results with SCUBA collections of Orconectes virilis and Cambarus bartoni in an Ontario oligotrophic lake was done during October. The effects of four bait types (chicken, pork liver, fish, and dry dog food), water temperature, and lunar phase on number and size of crayfish caught was assessed. The mean carapace length of crayfish taken in traps was larger than crayfish caught by divers. The sex ratio was biased toward males for trap catches, but was essentially 1:1 for SCUBA results. Although the number of crayfish caught by each bait type was not different, the size of crayfish was largest for traps with chicken, decreasing for liver, fish, and then dog food. Low temperatures were shown to reduce trapability, and the lunar phase (or cloudiness) may also affect size of crayfish catch. Several combinations of interactions between these factors and variation in habitat were also evaluated. Sommer, T.R., and C.R. Goldman. 1983. The Crayfish Procambarus clarkii from California rice fields: Ecology, problems, and potential for harvest. Freshwater Crayfish 5:418-428. The life history and distribution of P. clarkii in California is presented in conjunction with a survey of its effects on the rice industry. This species may inhabit two-thirds of the State of California, and appears to have a negative impact on a majority of the rice growing operations. The timing of flooding rice fields is shifted from Louisiana studies and often results in a shorter growing season. Crayfish are reported to begin reproduction in April when fields are reflooded, and young are produced continuously through the summer. Fields are drained starting in late August, and crayfish must often rely on burrows for survival through the winter, as many fields are left dry until spring. Sommer, T.R. 1984. The biological response of the crayfish Procambarus clarkii to transplantation into California rice fields. Aquaculture, 41:373-384. Established populations of crayfish resulting from range extension in Southern California were studied over a two-year period. Comprehensive life history data was collected to evaluate physical conditions, reproduction, growth, density, and biomass of crayfish. Generally, timing of life cycle events for crayfish in California were similar to crayfish in Louisiana. However, differences in the hydrological cycle resulted in a shift in periods of activity, decreased juvenile growth rates and biomass, and lower densities of adults. In spite of suboptimal conditions, these populations may be reaching harvestable levels. Stein, R.A., and J.J. Magnuson. 1976. Behavioral response of crayfish to a fish predator. Ecology, Vol. 57 No. 4, pp. 751-761. The smaller the crayfish, the more vulnerable it is to fish predation; therefore, they tend to alter A-12 their behavior more dramatically than larger, adult crayfish. Walking, climbing, and feeding were supressed in small crayfish by the presence of a predator; however, burrowing and chelae display were increased in all size groups of crayfish. Adult males were less affected by the presence of a predator than adult females. This is due to the males' larger chelae which offer more protection. Smaller crayfish will also select substrates offering the greatest protection against fish predation. Taub, S.H. 1972. Exploitation of crayfish by largemouth bass in a small Ohio pond. Progressive Fish Culture 34(l):55-58. A pond with an established population of the burrowing crayfish (Cambarus diogenes) was stocked with largemouth bass fingerlings to assess the population dynamics of these two species in association. Crayfish densities, bass growth and food habits, and observations of plant growth were measured over a two-year period. Initial growth of fish was good but slowed during the second year. Incidence of crayfish in bass stomachs remained high (>40%) in spite of a severe reduction in crayfish densities. Observations of plants showed that macrophyte biomass and diversity increased as the crayfish population was reduced. Therefore, predation by fish on crayfish populations can be very effective and result in major biological shifts in community structure. Weingartner, D.L. 1982. A field-tested internal tag for crayfish (Decapoda, Crustaceana, 43 (2): 181-188. Astacidae). Crayfish tagging should: 1) not disrupt behavior, growth or longevity; 2) survive molting and the lifecycle of the crayfish; 3) be applicable to all life stages; 4) provide individual recognition; and 5) provide recognition without dissection. A new method of hypodermically injecting color-coded fishing line (3.5 mm long) for large crayfish (carapace length) 17.5 mm, or sewing thread (1.5 mm long) for smaller crayfish parallel to the midline of the abdomen near the fifth sternite, was found to be superior to conventional tagging methods. Wiernicki, C. 1984. Assimilation efficiency by Procambarus clarfdi fed Elodea (Egera densa) and its products of decomposition. Aquaculture, 36: 203-215. The ability of four size classes of crayfish to assimilate two sizes of Elodea fragments was examined in a compartmentalized tank. Comparison was also made between different ages (0, 15, 33, and 45 days) of plant material to evaluate the importance of microorganisms in diet. A carbon 14 radiotracer was used to differentiate plant material from microorganisms in the diets of crayfish. Aging improved assimilation efficiency, and microorganisms were especially important for smaller sizes of crayfish due to their enhanced ability to handle smaller fragments than adults. Witzig, J.F., J.W. Avult, Jr., and J.V. Huner. 1983. Crawfish, Procambarus clarkii, growth and dispersal in a small south Louisiana pond planted with rice, Oryza sativa. Freshwater Crayfish 5:331-343. Crayfish growth and distribution in a culture pond was examined after flooding with regard to water depth, vegetation density, pond area, and time after flooding. Unharvested rice Oryza sativa, served as cover and the major food source. Crayfish densities were higher in shallow water (0.6 to 0.8 m), dense vegetation ( > 646 g/m, dry wt.), and center areas. No preference for A-13 deep or shallow areas was seen approximately 200 days after flooding when vegetation had decomposed in both areas. With decreasing nutrient resources, crayfish biomass decreased. Supplemental high protein feed was recommended to sustain crayfish growth. The author notes that terrestrial vegetation decomposes slower than aquatic vegetation and would serve as a better supplement. A-14 Appendix B Appendix B. Habitat sampled by each trap during each introduction period. Aquatic plants = POPE (Potamogeton pectinatus) and NAMA (Najas Marina), terrestrial plant = BAGL (Baccharis glutinosa), emergent plant = cattails. Trap Intro 1 Intro 2 Intro 3 Intro 4 1-1 1-2 1-3 1-4 1-5 1-6 1-7 1-8 Rock Rock Rock Rock Rock Rock Rock Rock Rock Rock Gravel Gravel Gravel Gravel Gravel Gravel Rock Rock Gravel Gravel Gravel Gravel Gravel Gravel Rock Rock Rock Rock Gravel Gravel Gravel Gravel 2-1 2-2 2-3 2-4 2-5 2-6 2-7 2-8 Rock Gravel Mud Mud Mud Mud Mud Mud Rock Gravel Mud Mud Mud Mud Mud Mud Rock Gravel Mud Mud Mud Mud Mud Mud Rock Gravel Mud NAMA Mud Mud Mud Mud 3-1 3-2 3-3 3-4 3-5 3-6 3-7 3-8 Rock Rock Mud Rock Rock Rock Rock Rock Rock Rock POPE Rock Rock Rock Rock Rock Rock POPE POPE Rock Rock Rock Rock Rock Rock NAMA NAMA NAMA Rock Gravel ' Gravel NAMA 4-1 4-2 4-3 4-4 4-5 4-6 4-7 4-8 Rock Mud Mud Rock Rock Rock Rock Rock Rock POPE Rock Rock Rock Rock Rock Rock Rock POPE POPE Rock Rock Rock Rock Rock Rock NAMA NAMA Rock Rock Rock Rock Gravel (Continued) B-2 Appendix B. (Continued) Aquatic plants = POPE (Potamogeton pectinatus) and NAMA (Najas Marina), terrestrial plant = BAGL (Baccharis glutinosa), emergent plant = cattails. Trap Intro 1 Intro 2 Intro 3 Intro 4 5-1 5-2 5-3 5-4 5-5 5-6 5-7 5-8 Rock Gravel Gravel Gravel Gravel Gravel Rock Rock Gravel Gravel Gravel Gravel Gravel Gravel Rock Rock Rock Gravel Gravel Gravel Gravel Gravel Gravel Rock Gravel Gravel Gravel Gravel Gravel Rock Rock Rock 6-1 6-2 6-3 6-4 6-5 6-6 6-7 6-8 Mud Mud Mud Mud Mud Rock Rock Rock POPE Mud Mud Mud Mud Brush Gravel Gravel Gravel Gravel Gravel Gravel Gravel Gravel Gravel Gravel Gravel Gravel Gravel Gravel Gravel Rock Rock Rock 7-1 7-2 7-3 7-4 7-5 7-6 7-7 7-8 Rock Rock Rock Rock Rock Rock Rock Rock Rock Rock Rock Rock Rock Rock Rock Rock Rock Rock Rock Rock Rock Rock Rock Rock Rock Rock Rock Rock Rock Rock Rock ' Rock 8-1 8-2 8-3 8-4 8-5 8-6 8-7 8-8 BAGL BAGL Mud Mud Mud Mud Mud Mud BAGL POPE POPE POPE Mud Mud Mud Mud POPE Mud Mud Mud Mud Mud Mud Mud NAMA NAMA Mud Mud Mud Mud Mud Mud Continued B-3 Appendix B. (Continued) Aquatic plants = POPE (Potamogeton pectinatus) and NAMA (Najas Marina), terrestrial plant = BAGL (Baccharis glutinosa), emergent plant = cattails (cont.). Trap Intro 1 Intro 2 Intro 3 Intro 4 9-1 9-2 9-3 9-4 9-5 9-6 9-7 9-8 Cattails Mud Mud Mud Mud Mud Mud Mud Cattails POPE Mud Mud Mud Mud Mud Mud Cattails Mud Mud Mud Mud Mud Mud Mud NAMA NAMA Mud Mud Mud Mud Mud Mud 10-1 10-2 10-3 10-4 10-5 10-6 10-7 10-8 Rock Rock Mud Mud Mud Mud Mud Rock Rock Rock Mud Mud Mud Mud Mud Mud Rock Mud Mud Mud Mud Mud Rock Rock Rock Rock Mud Mud Mud Rock Rock Rock 11-1 11-2 11-3 11-4 11-5 11-6 11-7 11-8 Trapline Established on 2/1/89 Rock Gravel Gravel Rock Mud Mud Brush Brush Rock Rock Gravel Gravel Rock Mud Brush Brush Rock Gravel POPE Rock Mud Brush Brush POPE Rock Gravel Gravel Gravel Mud Mud Rock Rock 12-1 12-2 12-3 12-4 12-5 12-6 12-7 12-8 Trapline Established on 2/1/89 Rock Rock Gravel Gravel Gravel Mud Mud Mud Rock Rock Gravel Gravel Gravel POPE Mud Mud Rock Gravel Gravel POPE POPE POPE POPE POPE Rock Gravel Gravel Gravel Mud NAMA ,NAMA NAMA (Continued) B-4 Appendix B. (Concluded) Aquatic plants = POPE (Potamogeton pectinatus) and NAMA (Najas Marina), terrestrial plant = BAGL (Baccharis glutinosa), emergent plant = cattails (cont.). Trap Intro 1 Intro 2 Intro 3 Intro 4 13-1 13-2 13-3 13-4 13-5 13-6 13-6 13-8 Trapline Established on 2/1/89 Rock Rock Gravel Gravel Gravel Brush Gravel Gravel Trapline Established on 5/2/89 Rock Rock Gravel Gravel Rock Brush Gravel Rock Rock Rock Gravel Gravel Gravel Brush Rock Rock Rock Rock Gravel Gravel Gravel Rock Gravel Gravel 14-1 14-2 14-3 14-4 14-5 14-6 14-7 14-8 Trapline Established on 2/1 1/89 Gravel Gravel Gravel Gravel Rock Rock Rock Rock Trapline Established on 5/2/89 Gravel Gravel Gravel Gravel Rock Rock Rock Rock Gravel Gravel Gravel Gravel Rock Rock Rock Rock Gravel Gravel Gravel Rock Rock Rock Rock Rock 15-1 15-2 15-3 15-4 15-5 15-6 15-7 15-8 Trapline Established on 2/1 1/89 Gravel Gravel Gravel Gravel Gravel Gravel Gravel Gravel Trapline Established on 5/2/89 Gravel Gravel Gravel Gravel Gravel Rock Rock Rock Gravel Gravel Gravel Gravel Gravel Rock Rock Rock Gravel Gravel Gravel Gravel Gravel Gravel Rock Rock B-5
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