Aquatic Insects in Montezuma Well, Arizona, USA: a Travertine Spring Mound with High Alkalinity and Dissolved Carbon Dioxide

Great Basin Naturalist

Volume 49 Number 1 Article 7

1-31-1989

Aquatic insects in Montezuma Well, Arizona, USA: a travertine spring mound with high alkalinity and dissolved carbon dioxide

Dean W. Blinn Northern Arizona University, Flagstaff

Milton W. Sanderson Northern Arizona University, Flagstaff

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Recommended Citation Blinn, Dean W. and Sanderson, Milton W. (1989) "Aquatic insects in Montezuma Well, Arizona, USA: a travertine spring mound with high alkalinity and dissolved carbon dioxide," Great Basin Naturalist: Vol. 49 : No. 1 , Article 7. Available at: https://scholarsarchive.byu.edu/gbn/vol49/iss1/7

This Article is brought to you for free and open access by the Western North American Naturalist Publications at BYU ScholarsArchive. It has been accepted for inclusion in Great Basin Naturalist by an authorized editor of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. AQUATIC INSECTS IN MONTEZUMA WELL, ARIZONA, USA: A TRAVERTINE SPRING MOUND WITH HIGH ALKALINITY AND DISSOLVED CARBON DIOXIDE

Dean W. Blinn 1 and Milton W. Sanderson 1

Abstract. —An annotated list ofaquatic insects from the high carbonate system of Montezuma Well, Arizona, USA, is presented for collections taken during 1976-1986. Fifty-seven taxa in 16 families are reported, including new distribution records for Arizona (Anacaena signaticollis, Laccobius ellipticus, and Crenitulus sp. [nr. debilis]) and the USA (Enochrus sharpi). Larval stages for Trichoptera, Lepidoptera, Megaloptera, Neuroptera, Chironomidae, and Anisoptera were absent even though the habitat lacks fish, and water temperature, dissolved oxygen, available food, and substrata appear adequate in Montezuma Well. The potential importance of alkalinity in restricting these insect groups is discussed.

Previous collections from the near-constant vents at the bottom of the pelagic zone and environment of Montezuma Well, Arizona, exits (4, 163 1 min ) through a side wall cavern have yielded several endemic species of plant in the travertine deposits (Cole and Barry and animal taxa (Polhemus 1976, Cole and 1973). The water level remains constant Watkins 1977, Czarnecki and Blinn 1979, throughout the year, and water temperature Landye 1981, Davies et al. 1985). Therefore, never varies more than ± 4.0 C, with an we believe that a thorough survey of the annual mean water temperature of 21.1 C aquatic insects occupying this high carbonate (Boucher et al. 1984). Concentrations of dis-

' system was warranted. To date, there have solved oxygen are greater than 6.0 mg 1 in been only a few published reports on the the littoral vegetation over a diel period. aquatic insects in Montezuma Well. Cole There are no fish in Montezuma Well, appar- (1965) prepared a list of aquatic insect species ently due to high concentrations of dissolved in the Well, and Polhemus (1976) described a CO,' (Cole 1983). The high alkalinity of the new heteropteran species (Ranatra mon- artesian water maintains a constant pH (6.5, tezuma) in the Well; Blinn et al. (1982) dis- s.d. ± 0.02) and a moderately high specific 2 cussed the nocturnal planktonic behavior of conductance (925-1,394 u,S cm" , 25 C). Ad- the endemic Ranatra species. Recently, Pol- ditional physical-chemical information on the hemus and Sanderson (1987) reported Mi- Well may be found in Cole and Barry (1973) crovelia rasilis from the Well, which was a and Boucher et al. (1984). new distribution record for the USA. Montezuma Well is an active, collapsed Methods travertine spring mound in the upper Sonoran Desert grassland of Arizona that has consider- Seasonal collections were initiated during ably higher concentrations of dissolved CO, 1976 and continued through 1986. Samples 1 T (>550 mg I" ) and alkalinity (>600 mg 1 were taken during both day and night with net

CaC03 ) than other aquatic habitats in the re- tows, grabs, and bottom dredges. The abun- gion. The system encloses an area of 0.76 ha dance of dominant insect taxa was estimated and has a mean depth of 6. 7 m (Cole and Barry from harvests of macrophytes taken from cir-

1973). The littoral zone supports a dense stand cular quadrats (573 cm ) or vertical 1-m tows of Potamogeton illinoensis Morong; a precipi- with a plankton net (153 |xm mesh). Ultra- tous drop beyond the submerged vegetation violet light traps were also employed dur- delineates a well-defined pelagic region (0.33 ing night periods to obtain flying adults. ha, maximum depth 17 m). Warm (24 C) water Assistance with species identification and/or enters through three or four artesian spring verification was provided by the following

Department of Biological Sciences, Northern Arizona University. Flagstaff, Arizona 86011

85 86 Great Basin Naturalist Vol. 49, No. 1 specialists: F. N. Young (Dytiscidae), H. P. doptera, Megaloptera, Neuroptera, Plecop- Brown (Elmidae), W. U. Brigham (Halipli- tera, and Anisoptera (Odonata) in our collections from Montezuma Well. In addition to dae), P. J. Spangler (Hydrophilidae), W. W. Wirth and VV. L. Grogan, Jr. (Ceratopogo- these groups, Chironomidae larvae were also nidae), B. V. Peterson (Diptera), J. Polhemus absent, but Cole (1965) reported the occur- (Veliidae), D. Bloodgood (Ephemeropta), and rence of Pentaneura sp. Furthermore, Cal- D. G. Huggins (Odonata). libaetis (Ephemeroptera) larvae were present in only one collection during July 1986. The absence of Plecoptera is not surprising Besults and Discussion because of their strict requirements for clean, Fifty-seven taxa of aquatic insects in 16 cool running waters (Harper and Stewart families and 5 orders were collected in Mon- 1984); however, the absence of Trichoptera, tezuma Well during this study (Table 1). Lepidoptera, Megaloptera, and Anisoptera Seven additional taxa have been reported by and the limited collections of Chironomidae Cole (1965). We assumed the presence of and Ephemeroptera are notable since larval larval stages indicated that insects reproduced stages for these aquatic insects are commonly in the Well. associated with hydrophytes and/or soft sedi- Our collections reported three new records ments within lakes and ponds (Brigham et al. of Coleoptera for the state of Arizona, includ- 1982, Merritt and Cummins 1984). The lack of ing Anacaena signaticollis, Crenitidus sp. (nr. predaceous fish, the abundance of potential debilis), and Laccobius ellipticus, and the first prey (Blinn et al. 1987), the near-constant records of Enochrus sharpi in the United annual water temperature (21.1 ± 4 C), the States. In addition, undescribed species of adequate dissolved oxygen concentrations 1 adult Brachypogon, Ceratoculicoides, Dasij- (6-14 mg l ), and the extensive vegetative helea, and Trichomyia were collected (per- refuge along the shore of Montezuma Well sonal communication, W. L. Grogan and would appear to provide a suitable habitat for B. V. Peterson) in light traps near the shore of all of these insect groups. Adults of Anisoptera Montezuma Well. have been observed flying over Montezuma The biting midge family, Ceratopogonidae, Well and ovipositing, but no nymphs have had the greatest diversity of species with 30 been collected. taxa, while the Hydrophilidae (Coleoptera) The absence of Trichoptera, Lepidoptera, were the next most diverse group (7 taxa plus Megaloptera, Neuroptera, and Anisoptera, as Hydrophilus reported by Cole 1965). The soft well as the infrequent occurrence of Chirono- bottom muds and abundant plant material in midae and Ephemeroptera (Callibaetis) in Montezuma Well may have provided a variety Montezuma Well, suggests that high concen- 1" of suitable substrata for members of the biting trations of dissolved C0 2 (550 mg ) and/or 1 midge family. Although not represented by alkalinity (600 mg l" CaC03 ) restrict hatching many individual taxa, the Nepidae (Ranatra and/or larval development by individuals in montezuma ), Coenagrionidae (Telebasis salna ), these aquatic insect groups, because other and Mesoveliidae (Mesovelia mulsanti) were physicochemical conditions in the Well ap- numerically the most abundant aquatic in- pear to be suitable for occupation. This agrees sects in Montezuma Well. Collections in the with the findings of Winget and Mangum top 20 cm of the vegetation yielded 87 nymphs (1979), who reported a significant negative 2 m~ (s.e. ± 21) of R. montezuma and 57.4 correlation between alkalinity and number 2 nymphs m (s.e. ± 19.8) of Mesovelia mul- of macroinvertebrate taxa in aquatic eco- santi during the summer, while integrated systems. collections in a 1-m water column in the vege- One hypothesis for the restriction of these 3 tation yielded 2,119 nymphs m (s.e. ± 372) insect groups is that the high carbonate alka- of Telebasis salna from June through August. linity in Montezuma Well may interfere with The absence of major aquatic insect groups cutaneous respiratory activities of the imma- in Montezuma Well may be of more ecological ture stages by forming deposits of CaC03 on interest than the actual occurrence of insect the body surface due to a shift in the carbonate species reported in Table 1. There were no equilibrium (Cole 1983). These deposits could larval representatives of Trichoptera, Lepi- greatly reduce the cutaneous surface area January 1989 BLINN, SANDERSON: MONTEZUMA WELL INSECTS 87

Table 1. Annotated list of aquatic insects from Mon- Table 1 continued. tezuma Well, Arizona, USA: * indicates new Arizona record; ** indicates new USA record; *** indicates Dasyhelea fasciigera Kieffer record reported only by G. A. Cole (1965). All identifica- Dasyhelea grisea group (Coquillett) tions for Ceratopogonidae were made from adults, and Dasyhelea messersmithi Waugh & Wirth larval stages were assumed to be present in Montezuma Dasyhelea mutabilis (Coquillett) Well. Dasyhelea pollinosa Wirth Dasyhelea pritchardi Wirth COLEOPTERA Dasyhelea sp. Forcipomyia brevipennis (Macquart) Dytiscidae Leptoconops (L.) torrens (Townsend) Celina occidentals ? (Young) Palpomyia occidentalis Grogan & Wirth Cybister ellipticus LeConte Parabezzia biennis (Coquillett) Desmopachria (Pachriodesma) mexicana Sharp Parabezzia sp. Neoclypeodytes cinctellus (LeConte) Phaenobezzia fulvithorax (Malloch) Thermonectes marmoratns (Hope) Stilobezzia antennalis (Coquillett) Elmidae Stilobezzia fuscula Wirth Microcylloepus similis (Horn) Stilobezzia pruinosa Wirth

Haliplidae Chironomidae Peltodytes dispersus Roberts ***Pentaneura sp. Peltodytes simplex (LeConte) Culicidae Hydraenidae ***Anopheles freeborni Aitken Ochthebius puncticollis LeConte Psychodidae Hydrophilidae Trichomyia sp. *Anacaena signaticollis (Fall) Stratiomyiidae Chaetarthria sp. ***Auparyphus sp. *Crenitulus sp. (nr. debilis Sharp) ***Odontomyia sp. **Enochrus sharpi Gundersen ***Hydrophilus sp. Heteroptera *Laccobius ellipticus (LeConte) Tropisternus columbianus Brown Belostomatidae ***Abedus breviceps Stal Hydroscaphidae Belostoma bakeri Montandon ***Hydroscapha natans LeConte Corixidae Ephemeroptera Cenocorixa wileyae (Hungerford)

Baetidae Hydrometridae Callibaetis sp. Hydrometra aemula Drake

Mesoveliidae Diptera Mesovelia mulsanti White Ceratopogonidae Naucoridae Alluaudomyia needhami Thomsen Ambrysus woodburyi Usinger Atrichopogon occidentalis Wirth Atrichopogon transversus Wirth Nepidae Atrichopogon sp. Banatra montezuina Polhemus Bessia sandersoni Grogan & Wirth ***Banatra quadridentata Stal Brachypogon sp. Veliidae Ceratoculicoides sp. Microvelia mulsanti White Culicoides brookmani Wirth Microvelia rasilis Drake Culicoides butleri Wirth & Hubert Culicoides haematopatus Malloch Odonata Culicoides salihi Khalaf Culicoides variipennis (Coquillett) Coenagrionidae Dasyhelea ancora (Coquillett) Enallagma civile (Hagen) Dasyhelea cincta group (Coquillett) Telebasis salna (Hagen)

available for gas exchange. It has been re- 1977). Those insect larvae that rely almost ported that many aquatic insect larvae, partic- exclusively on cutaneous respiration but lack ularly the trichopterans, rely exclusively on supplemental tracheal gills or the ability to the rich tracheal network located just beneath transport air stores may be unable to meet the the thin cuticle for gas exchange (Wiggins metabolic demands for oxygen in the high Great Basin Naturalist Vol. 49, No. 1

carbonate waters of Montezuma Well. It has Cole, G A , and W T Barry 1973. Montezuma Well, Arizona, as a habitat. Arizona Acad. Sci. 8: 7-13. also been proposed that the tracheal gills are J.

Cole, G A , and R. L. Watkins 1977. Hyalella mon- highly susceptible to damage by environmen- tezuma, a new species (Crustacea: Amphipoda) tal extremes since they serve as active uptake from Montezuma Well, Arizona. Hydrobiologia sites (Eriksen et al. 1984). The constant low 52: 175-184. pH (6.5) may also restrict the above described Czarnecki, D B , and D W Blinn 1979. Observations on southwestern diatoms. 2. Caloneis latiuscula insect groups from Montezuma Well. var. reimeri n. var., Cyclotella pseudostelligera f.

parva n. f. and Gomphonema montezumense n. Acknowledgments sp., new taxa from Montezuma Well National Monument. Trans. Amer. Microsc. Soc. 98: This study was supported in part by funds 110-114. provided to DWB from the Whitehall Foun- Davies, R W , R N. Sinchal, and D W Blinn 1985. Erpobdella montezuma, a new species of fresh- dation, Inc., and an Organized Research water leech from North America (Arizona, USA). Grant from Northern Arizona University. We Canadian J. Zool. 63: 965-969. thank Clay Runck for assistance with the col- Eriksen, C H , V H Resh, S S Balling, and G A Lam- lection and quantification of dominant insect berti Aquatic insect respiration. Pages 27-37 in species and the personnel at Montezuma R. W. Merritt and K. W. Cummins, eds., An introduction to the aquatic insects of North Amer- Well, especially Jack Beckman and Jim Cole- ica. 2d ed. Kendall/Hunt Publ. Co. 722 pp. man, for their excellent cooperation during Harper. P P., and K W Stewart 1984. Plecoptera. this study. Pages 182-230 in R. W. Merritt and K. W. Cum- mins, eds., An introduction to the aquatic insects of North America. 2d ed. Kendall/Hunt Publ. Co. Literature Cited 722 pp.

Landye, J J 1981. Current status of endangered, threat- Blinn, D W . R W Davies, and B Dehdashti. 1987. ened, and/or rare mollusks of New Mexico and Specialized pelagic feeding by Erpobdella mon- Arizona. U.S. Fish and Wildl. Serv., Office of tezuma (Hirudinea). Holarctic Ecology 10: Rare and Endangered Species, Albuquerque, 235-240. New Mexico. 35 pp. Blinn, D. W., C. Pinney, and M. W Sanderson 1982. Merritt, R W , and K W Cummins, eds 1984. An intro- Nocturnal planktonic behavior of Ranatra mon- duction to the aquatic insects of North America. 2d tezuma Polhemus (Nepidae: Hemiptera) in Mon- ed. Kendall/Hunt Publ. Co. 722 pp. tezuma Well, Arizona. J. Kansas Ent. Soc. 55: Polhemus, 1976. Notes on North American Nepidae 481-484. J. (Hemiptera: Heteroptera). Pan-Pacif. Ent. 52: Boucher, P., D W Blinn, and D B Johnson 1984. 204-208. Phytoplankton ecology in an unusually stable and W. Sanderson. 1987. Microvelia environment (Montezuma Well, Arizona, USA). Polhemus, J., M. Hydrobiologia 119: 149-160. rasilis Drake in Arizona: a species new to the United States (Heteroptera: Veliidae). Great Bricham, A R , W U Brigham, and A Gnilka, eds. 1982. Aquatic insects and oligochaetes of North Basin Nat. 47: 660. and South Carolina. Midwest Aquatic Enter- Wiggins, G. B. 1977. Larvae of the North American cad- prises, Mahomet, Illinois. disfly genera. University of Toronto Press, COLE, G A 1965. Final report to Montezuma Castle Toronto, Ontario. 401 pp. National Monument of investigations of Mon- WiNGET, R. N., AND F. A. Mangum 1979. Biotic condition tezuma Well. Report presented to National Park index: integrated biological, physical, and chemi- Service. 86 pp. cal stream parameters for management. U.S. De- 1983. Textbook of limnology. 3d ed. C. V. Mosby partment of Agriculture, Forest Service, Inter- Co., St. Louis, Missouri. 401 pp. mountain Region, Ogden, Utah. 51 pp.