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Contrasts Among Calanoid Copepods from Permanent and Temporary Ponds in Arizona

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Contrasts Among Calanoid Copepods from Permanent and Temporary Ponds in Arizona Contrasts among Calanoid Copepods from Permanent and Temporary Ponds in Arizona GERALD A. COLE • Reprinted from THE AMERICAN MIDLAND NATURALIST Vol. 76, No. 2, October, 1966, pp. 351-368 University of Notre Dame Press Notre Dame, Indiana Contrasts among Calanoid Copepods from Permanent and Temporary Ponds in Arizona' GERALD A. COLE Department of Zoology, Arizona State University, Tempe 85281 ABSTRACT: Thirty-seven plankton samples taken from Arizona ponds during October and November, 1962, contained calanoid copepods. Eighteen samples were judged to be from temporary waters, nineteen from permanent habitats. The species, in order of abundance, were Diaptomus albuquerquensis, D. siciloides, D. clavipes, D. novamexicanus, D. sanguineus and D. nudus. The first three occurred in both temporary and permanent waters; the last was present only in one permanent pond; and the other two species were restricted to temporary sites. Cephalothorax lengths for individual species were greater in tem- porary ponds than in permanent waters and, similarly, clutch sizes were greater. There was a strong correlation between female length and num- ber of eggs per clutch in D. clavipes, D. albuquerquensis and D. siciloides; the correlation was less in D. novamexicanus. Congeneric occurrence of adjacent-sized and similar-sized species is common in temporary ponds, but less so in permanent ponds. The reason for coexistence of similar- sized diaptomids, larger size, and more eggs per clutch in temporary ponds is either genetic, or the result of a better nutritional supply in ephemeral habitats. The trophic role of particles, which impart high turbidity to many temporary ponds, may be very important. In per- manent waters, predation could be a selective factor resulting in smaller individuals. INTRODUCTION In regions where aquatic habitats are abundant, certain calanoid copepod species are typical of permanent waters, while others occur principally in ephemeral pools. Near Louisville, Kentucky, for ex- ample, Diaptomus sanguineus Forbes, D. stagnalis Forbes, and Os- phranticum labronectum Forbes occur in vernal pools, whereas D. pallidus Herrick is characteristic of permanent lakes and ponds (Cole, 1953, 1959) . Farther north many more species occur, but a given species is rarely, if ever, found in both temporary and permanent habitats. Moreover, the temporary pools are usually vernal, clear and cool. In Arizona the situation is quite different. Water is relatively scarce; many ponds and lakes go dry at irregular intervals; temporary ponds are especially evident during the summer rainy season and are turbid and warm; and several calanoid species occur in both tem- porary and permanent habitats. Of the nine calanoids reported from Arizona, (Cole, 1961, 1963b, and present paper), at least four occur in both permanent and tem- porary waters in the state. This makes possible a study of the effects of differing conditions imposed on conspecific diaptomid populations in contrasting habitat types. Much of the present paper is concerned 1 This investigation was supported by Grant GB-154 from the National Science Foundation. 351 352 THE AMERICAN MIDLAND NATURALIST 76(2) with these effects. Interest in these and similar problems was aroused originally by an often-cited paper of Hutchinson (1951). The summer rains came late to central Arizona in 1962, not arriving until after mid-September. At Prescott the August precipi- tation was 6.4 cm below the standard normal, but September's rain was 4.3 cm greater than normal. Soon after the rains commenced, John G. Weise, then of this laboratory, began collecting phyllopod crustaceans from temporary ponds over a large area of the state. In addition he sampled many permanent, or at least more nearly per- manent ponds, and I did some concurrent collecting. Thirty-seven collections made between 12 October and 17 November included calanoid copepods. These were taken with a net of No. 20 bolting silk, and were qualitative only. The collections were preserved with 10% formalin. In the laboratory, calanoids were taken at random from the col- lections to establish relative numbers of different species, mean cephalothorax lengths, and other data discussed below. All collections were searched for uncommon species, and many were examined in their entirety, all copepods being counted and measured. In several instances, after adequate counts had been made, ovigerous females were examined selectively to determine mean clutch sizes. The results were many data concerning instantaneous samples of diaptomid populations from ponds which, with one exception, were visited only once. These data were examined for differences between inhabitants of temporary and permanent ponds. THE PONDS The bodies of water (Table 1) were classified as either permanent or temporary, despite the existence of intermediate types. The ponds of each category were symbolized and numbered chronologically so that those sampled on the earliest dates had the lowest numbers. Thus, P1 and P2 refer to two permanent ponds sampled on 12 Oct. 1962, the first collecting date; Ti and T2 refer to temporary ponds visited on the same day. In one instance, two collections were made from the same pond on different dates; these are treated here as two separate ponds, T6 and T7. Thirty-five ponds were either on the Colorado Plateau or at some- what lower elevations in the transitional mountainous region above the desert. Two ponds (P4 and P5), a few miles north of Congress Junction, Yavapai Co., at an elevation of 2,500 ft (760 m) , lay in surroundings showing Mohave Desert affinities. The 37 ponds ranged from about 610 to 2133 m above sea level. Water temperatures ranged from 24.4 C in October to 1.1 C in November. Most pH values at the time of collecting were high; none were above 9.0 or below 7.6. Transparencies, expressed as standard Secchi disc values, ranged from 2.5 cm in small turbid pools to 1.2 m in the permanent Holiday Lake (P19). Many ponds were either obviously permanent, or clearly of an 1966 COLE: CALANOID COPEPODS IN ARIZONA 353 ephemeral nature. Others were not easily classified. Unfortunately, the presence of anostracan, conchostracan, or notostracan crustaceans does not permit the automatic assigning of a pond to the temporary category. In the arid Southwest many permanent bodies of water show extreme surface-level fluctuations, and lack fish faunas. In such ponds phyllopod crustaceans sometimes appear, especially when dry margins become reflooded. These ponds probably satisfy the require- ments of phyllopods by having large temporary portions and by lack- TABLE 1.-Data on Arizona ponds from which calanoid copepods were collected, October-November, 1962. T = temporary; P = permanent a) i.- z r9 T 1 Yavapai 12 Oct 5300 18.9 8.6 3 T 2 Yavapai 12 Oct. 5300 24.4 8.7 2 T 3 Yavapai 27 Oct 5000 15.6 8.7 1 T 4 Yavapai 27 Oct 5000 21.1 8.8 1 T 5 Yavapai 27 Oct 5200 14.4 7.6 2 T 6 Yavapai 27 Oct 5000 15.6 7.8 3 T 7 Yavapai 3 Nov. 5000 15.0 .... 2 T 8 Coconino 3 Nov 6000 12.2 .... 2 T 9 Yavapai 8 Nov 5200 3.3 8.6 1 T 10 Yavapai 8 Nov 5300 12.2 .... 3 T 11 Yavapai 8 Nov 5000 15.6 8.4 3 T 12 Yavapai 10 Nov 5500 10.0 8.4 1 T 13 Coconino 16 Nov 5000 .... 3 T 14 Coconino 16 Nov. 5000 6.7 8.6 1 T 15 Coconino 16 Nov 5000 7.7 8.6 1 T 16 Navajo 17 Nov 5650 1.1 8.2 1 T 17 Navajo 17 Nov. 5975 2.2 8.4 1 T 18 Navajo 17 Nov 7000 2.2 8.2 4 P 1 Yavapai 12 Oct 5200 18.9 9.0 1 P 2 Yavapai 12 Oct 5200 18.9 9.0 2 P 3 Gila 21 Oct 5800 15.0 8.1 1 P 4 Yavapai 26 Oct 2500 20.0 8.7 2 P 5 Yavapai 26 Oct 2500 20.6 8.0 2 P 6 Yavapai 26 Oct 5000 15.6 8.7 1 P 7 Yavapai 27 Oct 5000 22.2 8.6 1 P 8 Yavapai 27 Oct 5000 22.2 1 P 9 Yavapai 27 Oct 5200 .___ 1 P 10 Maricopa 2 No v. 2000 18.9 8.4 1 P 11 Gila Nov 2000 20.0 7.6 1 P 12 Gila 2 NovNov 3000 18.9 8.4 2 P 13 Gila 4900 15.6 7.6 2 P 14 Maricopa 2 Nov. 3300 21.1 8.4 2 P 15 Yavapai 3 Nov. 4500 15.6 8.4 2 P 16 Coconino 3 Nov 6700 8.3 8.0 3 P 17 Yavapai 8 Nov. 5300 22.2 8.5 1 P 18 Yavapai 9 Nov 4900 16.7 7.8 1 P 19 Yavapai 10 Nov 5500 12.2 8.6 3 354 THE AMERICAN MIDLAND NATURALIST 76(2) ing vertebrate predators. Conversely, the presence of certain insects does not preclude classifying a pond as temporary. Mayfly naiads, notonectids, and other hemipterans are common in some temporary ponds. In general, ponds were considered permanent if they had obvious water sources such as springs or windmill pumps. In some cases information from people living nearby was useful in making decisions. Some stations are well-known permanent waters such as Kaibab Lake (P16), Holiday Lake (P19), Little Hell Canyon Reservoir (P17) and pond P13 near Payson, which was probably the "Small dammed reservoir. ." from which S. F. Light collected copepods during May, 1935 (see Wilson 1955:44). Several of the ponds observed later, during May 1963, still held water, while others formerly classified as temporary were dry at that time. Pond Ti in the Chino Valley was considered temporary although local residents said it had held water for several years prior to the summer of 1962.
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