Amphibians can be important consumers in both aquatic and terrestrial habitats and may represent an important energetic link between the two, particularly in the tropics, where amphibian species richness and abundance are high. In the past 20 years, amphibian populations have declined dramatically around the world; numbers have decreased catastrophically in protected upland sites throughout the neotropics, usually resulting in the disappearance of over 75% of amphibians at a given site, particularly those species that breed in streams. Most studies of amphibian declines have focused on identifying causes and documenting changes in adult abundance, rather than on their ecological consequences. Here, we review evidence for the potential ecological effects of catastrophic amphibian declines, focusing on neotropical highland streams, where impacts will likely be greatest. Evidence to date suggests that amphibian declines will have large‐scale and lasting ecosystem‐level effects, including changes in algal community structure and primary production, altered organic matter dynamics, changes in other consumers such as aquatic insects and riparian predators, and reduced energy transfers between streams and riparian habitats. Furthermore, because of habitat and functional differences between larvae and adults in most amphibians, the loss of a single species is akin to losing two species.
As part of the Tropical Amphibian Declines in Streams (TADS) project, we estimated macroinvertebrate production and constructed quantitative food webs for four headwater stream reaches in the Panamanian uplands: two that had experienced massive amphibian declines and two with unaffected amphibian populations. As expected for forested headwaters, allochthonous materials were the dominant energy source. Total macroinvertebrate biomass and production ranged from 231 to 360 mg ash-free dry mass m 22 and from 3.1 to 4.4 g ash-free dry mass m 22 yr 21 , respectively, and did not appear influenced by the presence or absence of amphibians. However, macroinvertebrate functional structure differed between pre-and post-decline sites, with shredder production significantly higher in pre-decline sites and scrapers significantly higher in post-decline sites. Taxonomic differences between pre-and post-decline sites were also evident. There was a shift in scrapers from smaller-bodied taxa (e.g., Psephenus) in pre-decline sites to larger-bodied groups (e.g., Petrophila) in post-decline sites. Detrital pathways were dominant in these systems, with shredders and collectors accounting for most energy flow. However, scrapers were well-represented and they were food-limited in these systems, particularly in the presence of larval amphibians at pre-decline sites. Ecological effects of catastrophic amphibian declines ranged from subtle shifts in taxonomic composition and functional structure of remaining consumers to changes in the availability and relative importance of autochthonous energy sources.
Allochthonous inputs of detritus represent an important energy source for streams in forested regions, but dynamics of these materials are not well studied in neotropical headwater streams. As part of the tropical amphibian declines in streams (TADS) project, we quantified benthic organic matter standing stocks and organic seston dynamics in four Panamanian headwater streams, two with (pre-amphibian decline) and two without (post-decline) healthy amphibian assemblages. We also measured direct litterfall and lateral litter inputs in two of these streams. Continuous litterfall and monthly benthic samples were collected for 1 year, and seston was collected 1-3 times/month for 1 year at or near baseflow. Direct litterfall was similar between the two streams examined, ranging from 934-1,137 g DM m -2 y -1 . Lateral inputs were lower, ranging from 140-187 g DM m -1 y -1 . Dead leaves (57-60%), wood (24-29%), and green leaves (8-9%) contributed most to inputs, and total inputs were generally higher during the rainy season. Annual habitat-weighted benthic organic matter standing stocks ranged from 101-171 g AFDM m -2 across the four study reaches, with *4 9 higher values in pools compared to erosional habitats. Total benthic organic matter (BOM) values did not change appreciably with season, but coarse particulate organic matter (CPOM,[1 mm) generally decreased and very fine particulate organic matter (VFPOM, 1.6-250 lm) generally increased during the dry season. Average annual seston concentrations ranged from 0.2-0.6 mg AFDM l -1 (fine seston, \754 lm [250 lm) and 2.0-4.7 mg AFDM l -1 (very fine, \250 lm [1.6 lm), with very fine particles composing 85-92% of total seston. Quality of fine seston particles in the two reaches where tadpoles were present was significantly higher (lower C/N) than the two where tadpoles had been severely reduced (P = 0.0028), suggesting that ongoing amphibian declines in this region are negatively influencing the quality of particles exported from headwaters. Compared to forested streams in other regions, these systems receive relatively high amounts of allochthonous litter inputs but have low in-stream storage.
1. We quantified production and consumption of stream-dwelling tadpoles and insect grazers in a headwater stream in the Panamanian uplands for 2 years to assess their effects on basal resources and energy fluxes. At the onset of our study, this region had healthy, diverse amphibian populations, but a catastrophic disease-driven decline began in September 2004, which greatly reduced amphibian populations. 2. Insect grazer production was 348 mg ash-free dry mass (AFDM) m )2 year )1 during the first year of the study and increased slightly to 402 mg AFDM m )2 year )1 during the second year. 3. Prior to amphibian declines, resource consumption by grazers (tadpoles and insects) was estimated at 2.9 g AFDM m )2 year )1 of algal primary production, which was nearly twice the estimated amount available. Insect grazers alone accounted for c. 81% of total primary consumption. During the initial stages of the declines, consumption remained at c. 2.9 g AFDM m )2 year )1 , but only 35% of the available resource was being consumed and insect grazers accounted for c. 94% of total consumption. 4. Production and resource consumption of some insect grazers increased during the second year, as tadpoles declined, indicating a potential for functional redundancy in this system. However, other insect grazer taxa declined or did not respond to tadpole losses, suggesting a potential for facilitation between tadpoles and some insects; differential responses among taxa resulted in the lack of a response by insect grazers as a whole. 5. Our results suggest that before massive population declines, tadpoles exerted strong top-down control on algal production and interacted in a variety of ways with other primary consumers. 6. As amphibian populations continue to decline around the globe, changes in the structure and function of freshwater habitats should be expected. Although our study was focused on tropical headwater streams, our results suggest that these losses of consumer diversity could influence other aquatic systems as well and may even reach to adjacent terrestrial environments.
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