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Survey and Distribution of Invertebrates from Selected Springs of the Edwards Aquifer in Comal and Hays Counties, Texas Author(S): James R

Survey and Distribution of Invertebrates from Selected Springs of the Edwards Aquifer in Comal and Hays Counties, Texas Author(S): James R

Southwestern Association of Naturalists

Survey and Distribution of Invertebrates from Selected Springs of the in Comal and Hays Counties, Texas Author(s): James R. Gibson, Scott J. Harden and Joe N. Fries Source: The Southwestern Naturalist, Vol. 53, No. 1 (Mar., 2008), pp. 74-84 Published by: Southwestern Association of Naturalists Stable URL: https://www.jstor.org/stable/20424894 Accessed: 17-02-2020 20:47 UTC

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This content downloaded from 147.26.181.231 on Mon, 17 Feb 2020 20:47:01 UTC All use subject to https://about.jstor.org/terms THE SOUTHWESTERN NATURALIST 53(1):74-84 MARCH 2008

SURVEY AND DISTRIBUTION OF INVERTEBRATES FROM SELECTED SPRINGS OF THE EDWARDS AQUIFER IN COMAL AND HAYS COUNTIES, TEXAS

JAMES R. GIBSON,* ScoTJ. HARDEN, AND JOE N. FRIES

National Fish Hatchery and Technology Center, United States Fish and Wildlife Service, 500 East McCarty Lane, San Marcos, TX 78666 (JRG, JNF) 8019 Riata Drive, San Antonio, TX 78227 (SJH) *Correspondent: randy [email protected]

ABSTRACT-We surveyed four spring sites of the Edwards Aquifer to determine current distribution and abundance of spring and cave-adapted invertebrates, including three that are federally listed as endangered. We found 18 from 12 families of endangered and stygobiontic fauna, similar to other surveys completed 10 years prior, and recorded new localities for Tethysbaena texana, Cirolanides texensis, Lirceolus hardeni, Lirceolus smithii, longipes, Stygobromus flagellatus, and Artesia subterranea. We also report discovery of new species of Bogidiellidae (near Parabogidiella), Ingolfiella, and Almueyzothyas.

RESUMEN-Examinamos cuatro sitios de nacimento de aguas del acuifero Edwards para determinar las distribuciones y abundancias de invertebrados adaptados a manantiales y cuevas, incluyendo 3 especies que estan en peligro de extinci6n a nivel federal. Encontramos 18 especies de 12 familias de fauna estigobi6nticas parecidas a muestreos realizados hace 10 afnos. Reportamos nuevas localidades para Tethysbaena texana, Cirolanides texensis, Lirceolus hardeni, Lirceolus smithii, Stygobromus longipes, Stygobromus flagellatus y Artesia subterranea. Tambien reportamos el descubrimento de nuevas especias de Bogidiellidae (similar a Parabogidiella), Ingofiella y Almuerzothyas.

The limestone aquifers of the Edwards Plateau Edwards Aquifer are listed as endangered by support about one-half of all springs in Texas, the United States Fish and Wildlife Service including some of the largest (Brune, 1981). (1997)-Comal Springs riffle Heterelmis Bowles and Arsuffi (1993) identified 91 endemic comalensis, Peck's cave amphipod Stygobromus species in the permanent aquatic habitats of the pecki, and Comal Springs dryopid beetle Stygo Edwards Plateau with >40 described species of parnus comalensis. These aquifer-dependent spe stygobionts (Hershler and Longley, 1986r, Barr cies are subject to population reduction, or and Spangler, 1992). This aquifer is the major possible extinction, if water quality or quantity source of water for the rapidly growing city of is decreased due to drought or over-pumping. San Antonio. It also is important for agriculture, They only occur in Comal and Hueco springs in fisheries, and tourism. Water flowing down the Comal County and San Marcos and Fern Bank Guadalupe River supplied by the aquifer pro springs in Hays County, Texas (Fig. 1). vides important coastal habitat for estuarine Comal Springs, located in the city of New reliant species such as economically important Braunfels, is the largest spring system (mean shrimp, crabs, and fishes (Bowles and Arsuffi, discharge = 8,200 during 1927-2005; United 1993), as well as waterfowl that feed on these States Geological Survey, http://nwis.waterdata. species. Thus, it is important to maintain the usgs.gov/tx/nwis/discharge/, to 9,900 L/s dur quality and a sufficient quantity of water in the ingl882-1926; Brune, 1981) in Texas and the Edwards Aquifer. Periodic surveys of the stygo southwestern United States. Comal Springs is a biontic communities can be useful to detect spring complex with numerous discharge loca changes within the aquifer that might affect tions occurring along the 1,300-m reach forming those that rely on this resource. Three aquatic Landa Lake. Tracer testing performed by the invertebrates associated with springs of the Edwards Aquifer Authority (EAA) in 2002

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FIG. 1-Sites in south-central Texas surveyed for invertebrates by drift netting and hand sampling at spring openings at (1) Comal Springs, (2) Hueco Springs, and (3) Fern Bank Springs during spring and summer 2003. San Marcos Springs (4) was sampled using cotton-cloth lures during spring 2004 and 2006. In small map of Texas, rectangle indicates expanded region of named counties and dark polygon represents the artesian zone of the Edwards Aquifer.

indicates convergent, conduit-dominated contribution flow to the springs and naphthalene paths in the Comal Springs area. Groundwater detected at Hueco and Comal springs from a discharging from Comal Springs originates diesel inspill in January 2000 about 6.5 km SW New regions as far as 225 km to the west. Discharged Braunfels suggest regional contribution (G. groundwater ranges in age from hours to Schindel, many pers. comm.). The occurrence of some years. The spring complex has had continuous of the same species at both Comal and Hueco flow with an exception in 1956, when flow springs ceased also suggests a regional component of for about 6 months after 7 years of drought flow. (Brune, 1975, 1981). San Marcos Springs, located in the City of San Hueco Springs, located 5.1 km N ComalMarcos, is the second largest spring system Springs, is a much smaller spring system (mean (mean discharge = 4,100 during 1916-1955; discharge = 1,100 during 1924-1978; Brune, Brune, 1981, to 4,900 L/s during 1956-2005; 1981, to 1,800 L/s during 2002-2006; UnitedUnited States Geological Survey, http://nwis. States Geological Survey, http://nwis.waterdata. waterdata.usgs.gov/tx/nwis/discharge/) in Tex usgs.gov/tx/nwis/discharge/) issuing from as. Sanallu Marcos Springs is a complex of many vial gravel about 90-150 m west of the artesian Guada springs upwelling into Spring Lake, lupe River. Hueco Springs, like Comal Springs, forming the head of the San Marcos River has a local and regional component; tracer (Brune, tests 1981). Records indicate this spring by Ogden et al. (1986) showed some complex local has not ceased flowing. Water discharg

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ing from San Marcos Texas. Springs We alsohas comparedmultiple our results with those sources; southern flow of from C. B. theBarr San (in Antoniolitt.). area in Bexar County that bypasses Comal Springs and flow from the MATERIALS northern AND area METHODS-We of the collected invertebrates aquifer, south of the Barton using Springsdrift nets groundwa (0.45 by 0.30-m rectangular opening, 250 gm mesh) at Comal, Fern Bank, and Hueco ter divide near Kyle, Hays springs County. (Fig. Groundwater 1). From 2 May to 16 August 2003, we from the southern flow sampled paths discharges each site with from five drift nets twice in both the southern portion of springSpring and Lake summer while exceptwater Fern Bank Springs, which from the northern flow was paths sampled discharges with two from nets once in spring and twice in summer, totaling 46 samples. We deployed nets for the northern portion of the lake (G. Schindel, about 24 h and checked them every 4-6 h. Because so pers. comm.). few specimens were found at Fern Bank Springs during Fern Bank Springs is located the first 8 km sampling E Wimberley period, nets were left out for 67-74 h and 13 km NW San Marcos. and Achecked small oncecave dailystream during the last two sampling periods. Specimens of H. comalensis and S. pecki large empties into a pool before diffusely flowing over a enough to identify in the field were returned to the bluff and cascading into area the whereBlanco they River. were Many collected and all other inverte seeps emerge near the base brates of fromthis cliff.nets wereThese preserved in 70% ethanol for springs are influenced by later local study. recharge We calculated and may combined drift time (sum of be perennial with recorded hours netsdischarge were placed ranging at a site) for each spring system and drift rate (number collected/day) of each species. from 140 L/s on 31 May 1975 to 9 L/s on 1 May During March 2004 and March and April 2006, we 1978 (Brune, 1981; C. B. usedBarr, cotton-cloth in litt.). lures in San Marcos Springs to see if Heterelmis comalensis was H. comalensisdescribed wasby Bossestill present et after C. B. Barr (in litt.) al. (1988) and was known collected only the from only Comalknown specimen from this location Springs until C. B. Barr in (in1992. litt.) This collected method was one adapted from the mop head and also the gauze technique (Hershler and Longley, adult from San Marcos 1986a) Springs. for collecting cave snails and used by G. Cole comalensis and Stygobromus for collecting pecki were isopods described (Holsinger and Minckley, 1971). and originally known only We fromlodge cottonComal Springscloths, about handkerchief-sized, (Holsinger, 1967; Barr and inside Spangler, spring orifices. 1992), butOver 3-5 weeks, microorganisms (primarily fungi) grow on the cotton substrate, serving both are stygobiontic and difficult to collect. T. as a probable food source for some invertebrates. After L. Arsuffi (in litt.) recommended we remove the use cloths of drift and place them in a tray filled nets for sampling these withspecies water, because invertebrates of poor are easily picked off un results (only a few S. pecki harmed. from We Comal recommend Springs this passive collecting tech and no S. comalensis) with nique afor Hess the samplerendangered and , S. comalensis and H. comalensis, because it is less labor intensive, less handpicking. C. B. Barr (in litt.) collected many damaging to the , and non-destructive to the S. pecki from the headwaters habitat of than the methods Comal Riverpreviously employed. by placing drift nets atAdditionally, spring orifices we reviewed and the literature and collec recorded a range extension tion by records capturing of the a singlestygobiontic species found at specimen at Hueco Springs. Comal, In SeptemberHueco, and Fern2002, Bank springs. We determined the current distribution of these species using this and J. Krejca (unpublished specimensdata) collected collected S. orpecki identified by the authors. We (verified by SJH) from Pantheralso enlisted Canyon taxonomic Well (18 expertise of D. Bowles, J. m deep), about 110 m from Holsinger, Comal andSprings. I. Smith. C. B. Barr (in litt.) also caught a few S. comalensis by drift netting at Comal Springs RESULTS and AND at Fern DISCUSSION-We Bank collected 18 sty Springs (31.5 km NE gobionticComal Springs, and listed range species (Table 1), repre extension). Neither S. comalensis senting nor12 familiesS. pecki havein 5 orders. Total numbers been found from other andspring drift orifices rates ofor listedwells, and stygobiontic species suggesting that these species collected are restricted at each locality to the during the 2003 drift known locations and not survey distributed are in throughout Table 2. Species currently listed as the aquifer (United States endangered Fish and were Wildlife present at the localities sam Service, 1997). pled by C. B. Barr (in litt.). To determine current distributionEndangered Species-We and abun collected S. comalensis dance of endangered and at other Comal spring Springs and and cave Fern Bank Springs. During adapted invertebrates ina combinedthe Edwards drift Aquifer time of 451.6 h at Comal region, we surveyed several Springs, spring we sites collected in central two adults and three larvae,

This content downloaded from 147.26.181.231 on Mon, 17 Feb 2020 20:47:01 UTC All use subject to https://about.jstor.org/terms March 2008 Gibson et al.-Invertebrates from the Edwards Aquifer 77 which is similar to C. B. Barr pecki(in litt.) during who a 455.6-h combined drift effort (1.2/ collected one adult and four larvae day). during C. B. aBarr (in litt.) collected only one 397.5-h combined drift time. Additionally, specimen during we a 96-h combined drift effort collected a single larva by hand sampling (0.2/day). and an adult on a cotton-cloth lure placed We infound a small S. pecki in gravel, rocks, and spring opening. Subsequently, we collected associated debristhree (leaves, roots, and wood) more adults on cloth lures placed directly in inseeps or near of springs, seeps, and upwellings spring runs and along the western of shoreline Comal Springs ofand Landa Lake, but we did not Landa Lake in 2004. In 2005, we observed observe them three in surrounding habitats of the adults on pieces of rotting wood spring resting runs and onlake. Many S. pecki collected at upwellings on the bottom of Landa Comal SpringsLake. were At entirely bright orange. All Fern Bank Springs, we collected othera single stygobiontic larva amphipods appeared white or from a spring at the base of a hillside slightly during yellow, but a their gut would sometimes 304.8-h combined drift time of two appear drift yellow, nets orange, tan, or dark brown. All the compared to C. B. Barr (in litt.) stygobionticwho collected fauna of Hueco Springs, including S. one adult and 12 larvae from a pecki,single were smallwhite. Color of some amphipods, and hillside spring during a 131.5-h combined driftin general, often is dependent on time. We placed one net in a small diet, cavewhich usuallywhere includes carotenoids produced the largest spring (depth = 0.15 by plants, m; algae,mean and probably some bacteria current velocity = 0.36 m/s) emerged (Negre-Sadargues and et al., 2000; Fraser and Bram collected stygobionts other than Stygoparnus. ley, 2004; Gaillard et al., 2004). It is possible that However, the diffuse streams issuing the orangefrom coloration the found within the Comal spring pool did not produce any stygobionts. Springs population We is derived from food sources placed the second drift net over a (e.g.,small leaf springlitter) that were scarce where S. pecki that flowed out of the base of the inhabitsbluff Huecodirectly Springs. into the Blanco River. Here, we Overall, collected we retrieved two H. comalensis daily stygobionts, including a single from larva each spring of sampled S. with drift nets at Comal comalensis during a 140-h total Springs.drift Priortime. to ourA survey, Norris (2002) variety of trees and shrubs were growing collected invertebrates on top at Comal Springs using of this travertine bluff. The face of thethis same bluff drift netswas we used, placed in the center covered with bryophytes and ferns of andthe spring produced runs >100 m downstream from the stalactites that dripped ground water spring heads, directly and found H. comalensis (mostly into the river. Water percolates larvae)through to be much this higher in the drift (as much as bluff, potentially supplying aquatic 18 areas beetles/day). for the It is possible that drift of H. adult dryopids and damp terrestrial comalensis areas wasfor higher the in Norris' survey because he larvae. Also, microbial growth, potentially on roots captured and beetles drifting from all areas organic debris found within the bluff, of spring provides runs upstream a of the drift net, which potential food source. During this study, encompassed too many few springs along the margins as S. comalensis were collected to compare well as numerous relative upwellings throughout the abundance among samples and with streambeds. previous These springs and upwellings range studies in which more aggressive from techniques large, high-flow, cave-like orifices to small, were used. While specific habitat low-flow,requirements gravel, sand, or mud seepages. We found of S. comalensis are unknown, it is H. possiblecomalensis primarily that in or near spring outlets, the preferred habitat is difficult to butsample microhabitat or that preferences still are not known. it comprises a small area, or both, Bowleswhich et resultsal. (2003) collected H. comalensis by in this beetle rarely being collected. hand sampling the streambed within spring runs Similar to C. B. Barr (in litt.), we foundand found S. the pecki beetles to be randomly dispersed to be abundant in our drift-net throughoutsamples runs; from however, the sampling resolu Comal Springs. We collected 173 tion specimenswas 1 m2 and microhabitat preferences during a 451.6-h combined drift might effort not have (9.2/ been detectable at that level. day) and C. B. Barr (in litt.) collected We used the cotton-cloth-lure154 method to specimens during a 384-h combined collect drift >50 effort Heterelmis sp. (adults and larvae (9.6/day). At Hueco Springs, we collected combined) from23 S. upper reaches of San Marcos

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TABLE 1-Distribution of stygobiontic and listed species found at Comal, Hueco, and Fern Bank springs, Texas. All records without citations were specimens collected or identified by authors. Counties are in uppercase and all located in Texas. Superscripts are as follows: a = new county record; b = new locality record; and c = first known record.

Taxa Distribution Almuerzothyas n. sp. COMAL: Comal Springsc Artesia subterranea COMAL: Comal Springsa HAYS: TSU artesian well (Holsinger and Longley, 1980), Ezell's Caveb (collected byJ. Krejca) Cirolanides texensis COMAL: Comal and Hueco springsb; Panther Canyon Well (Krejca, 2005) southern Edwards Plateau region in Texas and northern Mexico (Krejca, 2005) Comaldessus stygius COMAL: Comal Springs HAYS: Fern Bank Springsa Haideoporus texanus COMAL: Comal Springs HAYS: TSU artesian well (Young and Longley, 1976) Heterelmis comalensis COMAL: Comal Springs HAYS: San Marcos Springs Ingolfiella n. sp. COMAL: Comal Springsc Lirceolus hardeni BLANCO, COMAL, HAYS, KENDALL, AND TRAVIS (Lewis and Bowman, 1996; Krejca, 2005) Lirceolus pilus BANDERA: Lost Maples State Park (Lewis and Bowman, 1996) COMAL: Comal and Hueco springsb, Preserve Cave (Krejca, 2005) HAYS: San Marcos Springsa MEDINA: Valdina Farms Sinkhole (Lewis and Bowman, 1996) Mexiweckelia hardeni BEXAR: San Antonio River alluviuma COMAL: Comal and Hueco springsa MEDINA: Hondo Creek alluvium (Holsinger, 1992) Parabogidiella? BEXAR: Verstraeten well no. 1 (Holsinger and Longley, 1980) COMAL: Comal Springsa HAYS: TSU artesian well (Holsinger and Longley, 1980) Seborgia relicta COMAL: Comal and Hueco springsa HAYS: TSU artesian well (Holsinger and Longley, 1980); Ezell's Caveb (collected byJ. Krejca) MEDINA: Hondo Creek alluvium (Holsinger, 1992) Stygobromus flagellatus COMAL: Comal Springsa HAYS: TSU artesian well; San Marcos Springs; Ezell's and Rattlesnake caves (Holsinger, 1966, 1967; Holsinger and Longley, 1980) TRAVIS: Barton Springsa (collected by D. Chamberlain) Stygobromus longipes HAYS: San Marcos Springsa KENDALL: Cave without-a-name (Holsinger, 1966) KENDALL/COMAL: Honey Creek Cave (Reddell, 1985) COMAL: Comal and Hueco springs; Panther Canyon Well (collected byJ. Krejca) Stygobromus russelli widespread in central Texas (Holsinger and Longley, 1980) Stygoparnus comalensis COMAL: Comal Springs HAYS: Fern Bank Springs Tethysbaena texana COMAL: Hueco springsb wells in BEXAR, HAYS, and UVALDE (Stock and Longley, 1981)

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TABLE 2-Total numbers and drift rates (number/day) of stygobionts and listed species collected at Comal, Hueco, and Fern Bank springs, Texas, 2 May-16 August 2003. Stygobromus spp. were immature and probably were S. pecki and S. russelli.

Comal Springs Hueco Springs Fern Bank Springs Taxa No. No./day No. No./day No. No./day Crustaceans Stygobromus pecki 173 9.2 23 1.2 Stygobromus russelli 371 19.7 91 4.8 14 1.1 Stygobromus spp. 120 6.4 42 2.2 14 1.1 Hadziidae Mexiweckelia hardeni 54 2.9 Sebidae Seborgia relicta 13 0.7 7 0.4 Bogidiellidae Artesia subterranea 6 0.3 Asellidae Lirceolus hardeni 14 0.7 439 23.1 9 0.7 Lirceolus near pilus 144 7.7 102 5.4 Cirolanides texensis 2 0.1 4 0.2 Thermosbaenacea Monodellidae Tethysbaena (=Monodella) texana 3 0.2 Coleoptera Comaldessus stygius 10 0.5 1 0.1 Haideoporus texanus 3 0.2 Stygoparnus comalensis 5 0.3 1 0.1 Elmidae Heterelmis comalensis 37 2.0

Springs (Spring Lake) in the area collected where during C. B. our survey (Table 2). Addition Barr (in litt.) had collected aally, single we obtainedadult, several new records and made 10 years prior. Of these specimens, discoveries 15 wereat each locality, with taxa including preserved and identified as H. comalensis beetles, crustaceans, by D. and a mite (Table 1). Bowles. We collected 28 adults Atand Comal larvae Springs, of we collected two species of Heterelmis in only three springs instygobiontic this same areabeetles other than S. comalensis; both during the subsequent survey of were 29 upwellings dytiscids and rare in our samples. These throughout Spring Lake. It appears included that Haideoporus H. texanus (three larvae), comalensis primarily occurs in the known headwaters only from of the artesian well in San Marcos Spring Lake. Additionally, we found (Young two andother Longley, 1976; Longley and Span species of elmid (Microcylloepus pusillus, gler, 1977)Stenelmis and spring runs of Comal Springs sexlineata) in eight springs sampled (Bowles from and theStanford, 1997), and Comaldessus headwaters to 180-m downstream. stygius (10 adults), known only from Comal Other Stygobionts-All other stygobiontic Springs (Spangler fauna and Barr, 1995). We also collected by C. B. Barr (in litt.) found also a single were adult of C. stygius in a drift-net

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sample from Fern Bankstreams Springs, and springs a range in Blanco, Comal, Hays, extension for this rare Kendall, species. and However, Travis counties to of central Texas confirm this record, additional (Lewis and collecting Bowman, should 1996; Krejca, 2005). We be performed. As is characteristic collected them of the at family, all sites (Table 2) and report both of these species thelikely first are records predators for this as species at San Marcos larvae. Larvae of C. andstygius Fern haveBank springs.yet to beLirceolus pilus has been collected and described. recorded Longley from and Spangler a spring system in Bandera (1977) presumed that larvae County of H. and texanus a sinkhole pupate in Medina Co., Texas along the margins of their (Lewis aquatic and habitat Bowman, and in 1996). Krejca (2005) trapped air spaces within collected the aquifer isopods because of the Lirceolus in dytiscids typically pupate Preserve in damp Cave, areas Comal out of County, and found them water. Adult dytiscids to typically be genetically frequent similar the to typical L. pilus. We surface to replenish their found air supplyother carriedpopulations in a in Comal County subelytral chamber. Because resembling these this beetles species, are Lirceolus near pilus, subterranean, they may which locate was air the spaces dominant within isopod at Comal Springs the aquifer for this purpose. (92%), Nearbut itsprings, was less these common at Hueco Springs air spaces may be maintained (19%). dueAt Santo the Marcos relative Springs, we collected L. stability groundwater level,hardeni possibly on cotton-cloth providing lures, and found L. pilus, habitat for all life stages and of Lirceolus these beetles. smithii Other in drift samples collected by than in Australia, stygobiontic personnel beetles of the are Edwards rare Aquifer Research and throughout the world, Dataand the Center presence and of Aquarena three Center. Lirceolus taxa (in two families and smithii three previously genera) of wassuch recorded only from the beetles in these spring systemsartesian is well remarkable. at Texas State University in San Thermosbaenaceans are Marcos in a primitive(Bowman andcrusta Longley, 1976). cean order found in groundwater Most Stygobromus and marine caught in this study were habitats primarily in small the Mediterranean (estimated as <5 and mm long). This may be Caribbean regions. However,because they Tethysbaena were dislodged easier than larger (=Monodella) texana occurs ones, theyin caves were and drifting wells for dispersal, or the of Hays, Bexar, and Uvalde population counties near (Stockspring andopenings was comprised Longley, 1981). We collected mostly three of smallspecimens individuals. at We recorded those Hueco Springs; the first that record were of too this small species to readily in identify to species Comal County. as Stygobromus spp., although they most likely We collected Cirolanides consisted texensis of at both Comal S. russelli and and S. pecki. The most Hueco springs. These arecommon the first amphipod records collected for at all sites was S. this species at both locations, russelli but(Table their 2). occurThis species is the most rence there is not surprising widespread because stygobiontic Krejca amphipod in Texas (2005) collected it in (Holsinger Panther Canyon and Longley, Well 1980). It is found in adjacent to Comal Springs karst prior habitats to our and survey. gravel stream beds, and likely This marine-relict isopod is more is believedcommon into shallowhave aquifer habitats than descended from ancestors deep in cavethe Gulfsystems of Mexico (Holsinger, 1967). At Hueco (Bowman, 1964) and hasSprings, been thefound largest in cavesnumbers of S. russelli (10/ and phreatic groundwaters day) were throughout from shallow the seeps and less (2/day) southern Edwards Plateau were from region large of upwellings. Texas. Also, more S. pecki Recently, Botosaneanu (1.2/day) and Iliffe than (2002) S. russelli record (0.2/day) were collect ed a subspecies in caves ed fromof northern the artesian Mexico. well. This suggests that S. Krejca (2005) used genetic pecki analysis inhabits to deeper show C. portions of the Hueco texensis had a pattern ofSpring evolution system that than followed does S. russelli. Holsinger subterranean hydrological (1967) connectionsdistinguished com the Jlagellatus group of pared to the freshwater-derived Stygobromus, Lirceolus which isopods includes S. pecki, as having that appear to have insularfollowed patterns surface of river speciation being greatly drainages. The genus Lirceolusrestricted occurs in ranges, in caves, and occupying deeper springs, and riverbed groundwater alluvium in nichesTexas andas compared to the more Mexico. Lirceolus hardeni common has been tenuis found group, in cave which includes S. russelli.

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Because S. pecki has been collected co-occuronly near and or rarely drifts out into San Marcos within spring systems, it is likely Springs. a phreatic Genetic comparisons of the different species and does not disperse well populationsin downstream and species of this genus might be gravel beds where S. russelli occurs. useful Similarly, to show SJH degree of isolation and connec assisting J. Krejca and P. Sprouse tivity (unpublished of habitats as well as assist in clarifying data) found S. pecki near the upwellings and ofnot the group. further downstream in the gravel We at discovered Hueco an apparent groundwater Springs during summer 2003. However, adapted S. russellipopulation of Crangonyx pseudogracilis was in both areas. Because of the phreatic at Hueco nature Springs. Some other amphipods in the of S. pecki and the hydrological same connection gracilis species group are stygomorphic and demonstrated by the aforementioned restricted diesel spill, to groundwater habitats (Zhang and we suspect this species inhabits deep Holsinger, groundwa 2003). The three specimens taken ter niches in the area around Comal from and HuecoHueco Springs have somewhat degenerate springs and follows hydrological eyes connections and little-to-no pigmentation. The same between these locations. species was in an interstitial habitat within Stygobromus longipes previously was stream-bank known only gravels of the Guadalupe River in from Cave Without-A-Name and Kendall Honey County,Creek about 50 km W Hueco Springs Cave in Kendall and Comal counties (pers. (Holsinger, observ., SJH). The specimens from Ken 1966, 1967; Reddell, 1985). We collected dall County a single resemble those from Hueco Springs specimen of S. longipes on the cotton-cloth in having lures reduced eyes and pigmentation. placed in San Marcos Springs, extending Gibson (2000) the found C. pseudogracilis in sandy known range into Hays County. springsStygobromus throughout eastern Texas with the flagellatus previously was known only typical from pigmentation Ezell's and well-developed eyes. Cave, Rattlesnake Cave, the artesian This well species at Texas is widespread in the east-central State University, and San Marcos Springs,United States all in and southern Canada, with a few the city of San Marcos (Holsinger, possibly 1966; introduced Hol populations in Arizona and singer and Longley, 1980). We identified Nevada, andspeci likely introduced populations in mens as S. flagellatus collected western by Dee Europe Ann (Zhang and Holsinger, 2003). Chamberlain (unpublished data) The in populations 2003 at in Comal and Kendall counties Barton Springs and JRG collected may represent a single the southwestern range limit for specimen in a drift survey of Comal the species. Springs extending the range of this species Holsinger from Hays and Longley (1980) considered County, north to Travis and south amphipods to inComal the family Crangonyctidae, includ counties. In 2005, we found S. longipes ing Stygobromus and and S. Crangonyx, to be of freshwa flagellatus in drift samples at San Marcoster origin. Springs All other stygobiontic amphipods collected by Edwards Aquifer Research collected andduring Data our survey are species that they Center and Aquarena Center, indicating considered to these be derived from relict marine populations are sympatric. Holsinger ancestors and that mightLong have colonized the region ley (1980) reported Stygobromus bifurcatus during regression (tenuis of a Cretaceous sea. group) from San Marcos Springs. We Thus, collected theresix Artesia subterranea at Comal have been four species of Stygobromus Springs during collected our survey extending the known here, the most recorded together at range any from locality. Hays County to Comal County. Within this genus, co-occurrence Because is rareA. subterranea and had previously been usually involves different species collectedgroups, in Ezell'swith Cave byJ. Krejca while diving hybridization being even rarer in(Holsinger, the aquifer during September 2006 (verified 1978). Although S. flagellatus is the byJRG) closest and fromknown the artesian well at Texas State relative to S. longipes (Holsinger, 1967),University we (59.5 found m deep) in San Marcos (Hol no obvious hybridization (intermediate singer and Longley, charac 1980), it is likely that this ters) in >100 specimens examined. species We primarily found inhabits deeper areas of the only a few S. longipes in samples aquifer. that Artesia were welbourni is the only other known dominated by hundreds of S. flagellatus species in thisand genus S. and is represented by three russelli. Presumably, S. longipes occupies specimens acollected differ from open water of a deep ent niche and area than S. flagellatus phreatic in orderpool within to Border Cave, Culberson

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Co., Texas (Holsinger, ingolfiellids1992). Another in the bogidiel subterranean freshwaters of lid was collected in the mainland drift at Comal North Springs America in and, perhaps like the November 2006 (pers. thermosbaenacean observ., JRG). TheTethysbaena two texana, their clos specimens likely are in est the known genus relativesParabogidiella may occur in the Mediterra and may be the same rarenean species region, mentioned appearing by to have a Tethys Sea Holsinger and Longley origin. (1980) This collected small inand the primitive suborder of well at Texas State University, amphipod Hays (Ingolfiellidea) County, and has been found in another well in Bexar interstitial County. marine, All of brackish,these and fresh ground specimens were missing water the habitats last pair worldwide of legs, (Stock, 1977; Ruffo and making identification inconclusive.Vonk, 2001). TheParabogidiella apparent rarity of this species americana, the only maydescribed be due species to its insmall the size and potential to genus, has been collected inhabit in thetight well interstitial at Texas spaces that are difficult State University, two wells to sample. in Bexar It County,also is possibleand that this species Honey Creek Cave inoccupies Comal the and interstitial Kendall groundwater of stream counties (Holsinger and beds Longley, similar to1980; those Fiers where M. hardeni and S. and Iliffe, 2000). These relicta specimens can occur. and A. subterranea are the only Holsinger bogidiellids and Longley reported (1980), suggested that north of Mexico. invasion of adjacent, interstitial habitats could We collected Mexiweckelia have hardeni occurred only in at theComal Tertiary and that the Springs during this study. Balcones This Fault species Zone was provides first a potential mixing discovered living in the zone interstitial for karst-groundwater alluvial ground fauna and those from water within the gravel the banks adjacent (hyporheic interstitial zone) of habitat of the Texas Hondo Creek in Medina coastal County plain. This(Holsinger, idea is supported by the fact 1992). It has since been that found some at Comalof the Springssame taxa (e.g., M. hardeni and (C. B. Barr, in litt.) and S. alluvial relicta deposits mentioned along earlier)the subsequently have San Antonio River, beenBexar found Co., Texasby SJH (pers. in alluvial biotopes in the observ., SJH). The other coastal two species plain andin this areas genus within the Balcones Fault occur in northern Mexico Zone. (HolsingerFurther evidence and Min that marine fauna may ckley, 1971; Holsinger, have 1973). invaded karst habitats through a hyporheic We collected Seborgia corridor relicta isat thatComal all andof the larger stygobionts Hueco springs, as did knownC. B. Barr from (in litt.).the Edwards This Aquifer, such as species was described from salamanders, the artesian fishes, well and at shrimps, are of ancient Texas State University freshwaterin San Marcos lineages. (Holsinger All of these animals also are and Longley, 1980) mobile,and later swimming found forms.in the The marine relicts, on hyporheos of Hondo theCreek contrary, with M.are hardeniall relatively small and benthic. (Holsinger, 1992). Seborgia We hershleri,collected manyfound nymphalin a and adult speci West Texas spring in Val mens Verde of County,a new species is a close of hydryphantid water relative to S. relicta (Holsinger, mite from 1992), a single and Pesce spring orifice at Comal and Iliffe (2002) collected Springs. S. near This hershleri mite, Almuerzothyas in a n. sp., belongs cave in Tamaulipas, Mexico. to a groupThe remaining of genera two distributed throughout members of this genus tropical are brackish and warm-temperate water, Old parts of the world World species-Seborgia and minima its closest isolated known in relative an is a species living in island lake in the South Costa Pacific Rica (Bousfield, (Goldschmidt 1970) and Gerecke, 2003; I. and Serborgia schieckei Smith,on the coast pers. of comm.). an island in the Indian Ocean (Ruffo, As apparent1983). by this and other surveys, the Comal Springs is the Edwards only localityAquifer forhas aan rich and diverse fauna undescribed species of that Ingolfiella, reflects itsrepresented complex geological structure and by a single specimen captured history. in Wethe recommenddrift during occasional surveys to 2004 (pers. observ., JRG). monitor Valentina subterranean Iannilli, S.communities for changes Ruffo, R. Vonk, and J. that Holsinger could reflect(pers. comm.) adverse conditions within the currently are describing aquifer this and and another to record species new faunal localities that from Bustamante, Mexico, hint toward that appear connectivity to be within the aquifer. closely related. These Additionally, are the only we suggest known use of genetic analysis

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on the more common taxa that Botosaneanu,could reflect L., and T. M. Iliffe. 2002. Notes on the hydrological connections, degree of isolationintraspecific of variability of Cirolanides texensis Bene cave and spring fauna, and geological dict, 1896 and (Isopoda: Cirolanidae) from Texas and taxonomic history, which are all important Mexico. Bulletin in de L'Institut Royal des Sciences understanding the complex nature Naturelles of the aqui de Belgique, Biologie 72:113-117. fer for conservation of the rare Bousfield, and endemic E. L. 1970. Terrestrial and aquatic amphi pod Crustacea from Renneil Island. Natural History species and associated habitats and management Museum of Rennell Island British Solomon Islands as a major water source. 6:155-168. Bowles, D. E., and T. L. Arsuffi. 1993. Karst aquatic Special thanks go to C. Thompson for help in all ecosystems of the Edwards Plateau region of central levels including finding funding, correspondence, Texas, USA: a consideration of their importance, reviewing, and field work. We thank the threats Keith to their and existence, and efforts for their Pfeuffer families for allowing us access to springs on conservation. Aquatic Conservation: Marine and their land. We also thank D. Bowles (National Park Freshwater Ecosystems 3:317-329. Service), J. Holsinger (Old Dominion Bowles,University), D. E., and R. and Stanford. 1997. A new distribu I. Smith (Agriculture and Agri-Food Canada) for their tional record for Haideoporus texanus (Cole?ptera: taxonomic expertise. We appreciate BIO-WEST, Inc., Dytiscidae), a stygobiontic beetle from the Edwards City of Austin Watershed Protection and Development Aquifer, Texas. Entomological News 108:297-299. Review Department, the Edwards Aquifer Bowles, Authority, D. E., C. B. Barr, and R. Stanford. 2003. and Zara Environmental Ltd. for providing us addi Habitat and pheneology of the endangered riffle tional information, as well as the staff at Aquarena beetle Heterelmis comalensis and a coexisting species, Center (notably E. Chappell and T. Kunz) and the Microcylloepus pusillus, (Cole?ptera: Elmidae) at Edwards Aquifer Research and Data Center for Comal Springs, Texas, USA. Archiv f?r Hydrobio supplying specimens. Thanks go to T. Castro, J. Gracia, logie 156:361-383. B. Grod,J. Hulbert, andJ. Thompston for their help in Bowman, T. E. 1964. Antrolana lira, a new genus and the field. We also thank M. Alexander, T. Arsuffi, D. species of troglobitic cirolanid isopod from Madi Bowles, T. Brandt, G. Longley, and J. Fowler-Propst for son Cave, Virginia. International Journal of Spele reviewing drafts. This project was a cooperative effort ology 1:231-236. between the San Marcos National Fish Hatchery and Bowman, T. E., and G. Longley. 1976. Redescription and Technology Center and Texas State University. Fund assignment to the new genus Lirceolus of the Texas ing, in part, was provided by a grant from Austin water slater, Asellus smithii (Ulrich) (Crustacea: Ecological Services, United States Fish and Wildlife Isopoda: Asellidae). Proceedings of the Biological Service. Endangered species collection operated under Society of Washington 88:489-496. Texas permit SPR-0390-045 (National Fish Hatchery Brune, G. 1975. Major and historical springs of Texas. and Technology Center), federal permit TE676811-9 Texas Water Development Board Report 189, (National Fish Hatchery and Technology Austin. Center), Texas permit SPR-0503-302 (Texas State University), Brune, G. 1981. Springs of Texas. Branch-Smith, Inc., and federal permit TE068895-0 (Texas StateFort Worth, Universi Texas. ty). A representative series of specimens were deposited Fiers, F., and T. M. Iliffe. 2000. Nitocrellopsis texanan. sp. in the Stephen F. Austin State University Invertebrate from central TX (USA) and N. ahaggarensis n. sp. Collection. Voucher specimens of Heterelmis comalensis from the central Algerian Sahara (Copepoda, from San Marcos Springs are in David Bowles' personal Harpacticoida). Hydrobiologia 418:81-97. reference collection at the National Park Service and Fraser, P. D., and P. M. Bramley. 2004. The biosynthesis deposited in the collection at Texas A&M and nutritional uses of carotenoids. Progress in University. Views presented do not necessarily reflect Lipid Research 43:228-265. those of the United States Fish and Wildlife Service. Gaillard, M., C. Juillet, F. Cezilly, and M. Perrot Minnot. 2004. Carotenoids of two freshwater Literature Cited amphipod species (Gammarus pulex and G. roeseli) and their common acanthocephalan parasite Poly Barr, C. B., and P. J. Spangler. 1992. A newmorphus genus minutus. and Comparative Biochemistry and species of stygobiontic dryopid beetle, PhysiologyStygoparnus Part B: Biochemistry and Molecular comalensis (Cole?ptera: Dryopidae), Biologyfrom 139:129-136. Comal Springs, Texas. Proceedings of the Gibson, Biological J. R. 2000. Macroarthropod communities of Society of Washington 105:40-54. sandy springs of East Texas. M.S. thesis, Texas A&M Bosse, L. S., D. W. Tuff, and H. P. Brown. University, 1988. College A new Station. species of Heterelmis from Texas Goldschmidt, (Cole?ptera: T., and R. Gerecke. 2003. Studies on Elmidae). Southwestern Naturalist 33:199-203. hydryphantid mites (Acari: Actinedida: Hydrachni

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dia) from Central and of deep-seaSouth shrimps (Crustacea,America. Decapoda, Alvino Pages 83-150 in An acarological carididae) tribute from a hydrothermal to David area of the MidR. Cook (from Yankee Springs toAtlantic Wheeny Ridge. Comparative Creek) Biochemistry and(I. M. Smith, editor). Indira PhysiologyPublishing Part A: Molecular House, and Integrative West Bloomfield, Michigan. Physiology 127:293-300. Hershler, R., and G. Longley. Norris, C.1986a. W. 2002. Effects Hadoceras of variable flows ontaylori, a new genus and species of invertebrate phreactic drift in Comal Springs, Hydrobiidae Texas. M.S. (Gastropoda: Rissoacea) from thesis, Southwest south-central Texas State University, San Marcos. Texas. Proceedings of the Biological Ogden, A. E., R. A.Society Quick, and S. R. Rothermel.of Washing 1986. ton 99:121-136. Hydrochemistry of the Comal, Hueco, and San Hershler, R., and G. Longley. Marcos springs, 1986& Edwards Aquifer, Phreatic Texas. Pages hydro biids (Gastropoda: Prosobranchia) 115-130 in The Balcones Escarpment:from geology, the Ed wards (Balcones Fault Zone) hydrology, Aquifer ecology and social region, development in south central Texas. Malacologia central 27:127-172. Texas (P. L. Abbott and C. M. Woodruff, Holsinger, J. R. 1966. Subterranean Jr., editor). Geological Society amphipods of America Annual of the genus Stygonectes (Gammaridae) Meeting, San Antonio, from Texas. Texas. Amer ican Midland Naturalist Pesce, 76:100-124. G. L., and T. M. Iliffe. 2002. New records of cave Holsinger, J. R. 1967. Systematics, dwelling mysids from the Bahamas speciation, and Mexico with and distribution of the subterranean description oiPalaumysis amphipodbahamensisn. sp. (Crustacea: genus Stygonectes (Gammaridae). Mysidacea). United Journal of Natural States History 36:265-278. National Museum Bulletin 259:1-176. Reddell, J. R. 1985. Cave biology of Honey Creek. Holsinger, J. R. 1973. TexasTwo Caver 30:133-134.new species of the subterranean amphipod Ruffo, genus S. 1983. Nuovi Mexiweckelia Anfipodi mesopsammici delle (Gam maridae) from Mexico and Isole Texas, Andamane (Crust. with Amphipoda) notes (Studi suion the origin and distribution of Crostacei the Anfipodi, genus. XCVIII). AssociationBollettino del Museo for Mexican Cave Studies Bulletin Civico Storia Naturale 5:1-12. di Verona 10:485-509. Holsinger, J. R. 1978. Systematics Ruffo, S., and R. Vonk. of 2001. theIngolfiella subterranean beatrids, new amphipod genus Stygobromus species (Amphipoda: (Crangonyctidae), Ingolfiellidae) from subterra part II: species of the easternnean waters of Slovenia. United Journal of States. Smithsonian Contributions Biology to21:484-491. Zoology 266:1-144. Holsinger, J. R. 1992. Four Spangler, new P. J., andspecies C. B. Barr. 1995. of A new subterranean genus and amphipod crustaceans (Artesiidae, species of stygobiontic dytiscid Hadziidae, beetle, Comaldessus Se bidae) from Texas, with stygius (Cole?ptera:comments Dytiscidae: Bidessini) on from their phylogenetic and biogeographic Comal Springs, Texas. relationships. Insecta Mundi 9:301-308. Tex as Memorial Museum, Stock,Speleological J. H. 1977. The zoogeography Monographs of the crustacean 3:1-22. suborder Ingolfiellidea, with descriptions of new West Indian taxa. Studies on the Fauna of Curacao Holsinger, J. R., and G. Longley. 1980. The subterra and Other Caribbean Islands 55:131-146. nean amphipod fauna of an artesian well in Texas. Smithsonian Contributions to Zoology 308:1-62. Stock, J. H., and G. Longley. 1981. The generic status Holsinger, J. R., and W. L. Minckley. 1971. A new genus and distribution of Monodella texana Maguire, the and 2 new species of subterranean amphipod only known North American thermosbaenacean. crustaceans (Gammaridae) from northern Mexico. Proceedings of the Biological Society of Washing ton 94:569-578. Proceedings of the Biological Society of Washing ton 83:425-444. United States Fish and Wildlife Service. 1997. Endan Krejca, J. K. 2005. Stygobite phlylogenetics as a tool for gered and threatened wildlife and plants; final rule determining aquifer evolution. Ph.D. dissertation, to list three aquatic invertebrates in Comal and University of Texas, Austin. Hays counties, TX, as endangered. Federal Register 62:66295-66304. Lewis, J. L., and T. E. Bowman. 1996. The subterranean Young, F. N., and G. Longley. 1976. A new subterranean asellids of Texas (Crustacea: Isopoda: Asellidae). Proceedings of the Biological Society of Washing aquatic beetle from Texas (Cole?ptera: Dytiscidae ton 109:482-500. Hydroporinae). Annals of the Entomological Soci Longley, G., and P. Spangler. 1977. The larva of a new ety of America 69:787-792. subterranean water beetle, Haideoporus texanus Zhang, J., andJ. R. Holsinger. 2003. Systematics of the (Cole?ptera: Dytiscidae: Hydroporinae). Proceed freshwater amphipod genus Crangonyx (Crangonyc ings of the Biological Society of Washington tidae) in North America. Virginia Museum of 90:532-535. Natural History Memoir 6:1-274. Negre-Sadargues, G., R. Castillo, and M. Segonzac. Submitted 16 January 2006. Accepted 25 June 2007. 2000. Carotenoid pigments and trophic behaviour Associate Editor was Joseph P. Shannon.

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