Research Article |
Corresponding author: David C. Houghton ( david.houghton@hillsdale.edu ) Academic editor: Ralph Holzenthal
© 2020 David C. Houghton, Ryan Lardner.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Houghton DC, Lardner R (2020) Ash-free dry mass values for northcentral USA caddisflies (Insecta, Trichoptera). ZooKeys 951: 37-46. https://doi.org/10.3897/zookeys.951.49790
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Ash-free dry mass (AFDM) values are presented for the adult stage of 63 caddisfly species commonly found throughout the northcentral US. Weights ranged from 0.01 mg for the smallest species to 7.22 mg for the largest. These values represent the first published data on the AFDM of the adult stage of Trichoptera, and can be used in other studies for more precise assessments of stream conditions without destruction of specimens. This increased precision is demonstrated herein by re-analyzing a previously published data set.
ash-free dry mass, biomass, caddisfly, Great Lakes, organic, Trichoptera
The organic biomass of organisms is one of the most important quantifiable variables in ecological studies. Measurements of biomass are informative about ecosystem production, metabolism, food web ecology, and the overall health and biotic integrity of the community (
There are several measurements used to express the biomass of organisms, including wet mass, dry mass, and ash-free dry mass (AFDM). To determine AFDM, specimens are incinerated at temperatures high enough to volatilize organic tissue but not inorganic tissue. The difference between pre-incineration and post-incineration weights reflects the mass of the organic tissue volatized. AFDM is considered the most accurate measurement of biomass since it encompasses the biologically active tissue (
Various parameters of immature aquatic insect assemblages, including their AFDM, have been used for many years to assess the functioning and biotic integrity of aquatic ecosystems. Some challenges to using the immature stage, such as the difficulty of sampling all aquatic microhabitats representatively and identifying specimens to the species level, can be alleviated by using the winged adult stage, particularly that of taxonomically and ecologically diverse groups such as the caddisflies (Trichoptera) (
Due to the necessity of maintaining museum collections of the taxonomically important caddisfly adults, most researchers are understandably reluctant to destroy them in order to obtain AFDM values. Indeed, while many studies have published data on caddisfly larvae (
We have been collecting caddisfly adults in the northcentral US since 2000, mostly utilizing an 8-watt ultraviolet light placed over a white pan filled with 80% EtOH. Such devices can capture 1000s of specimens during a single evening of heavy flight activity. Collected specimens are preserved in 80% EtOH for long-term storage, which limits decomposition and loss of organic biomass over time (
Species were chosen for biomass determination largely due to practical considerations. The weight of single specimens of most species is lower than the detection limit of most standard balances. Thus, specimens needed to be weighed in groups of 5 to 500 depending on the size of the species. This limitation meant that we could only determine biomass for abundant species for which we had ample extra specimens. Likewise, the specimen collecting localities that we chose were simply the ones with the most available specimens. Most of these specimens were from Michigan, with some from Indiana, Minnesota, and Wisconsin (Figure
To determine organic biomass, specimens of each tested species were taken from their vials of EtOH and placed into pre-dried porcelain crucibles. Crucibles containing the specimens were dried at low heat over a hot plate for several h until all of the EtOH had evaporated and the specimens appeared completely dry. The crucibles and specimens were then further dried for 2 h at 60 °C in a drying oven and then slowly cooled to room temperature before weighing. Crucibles and specimens were then transferred to a muffle furnace and incinerated at 500 °C for 3 h. After cooling to room temperature in the muffle furnace, the resulting material was transferred back to the drying oven, dried for 1 h at 60 °C, cooled to back room temperature, and weighed. AFDM was calculated as the final mass of material remaining after incineration subtracted from the mass of specimens before entering the muffle furnace. Total AFDM per sample divided by the number of specimens in that sample calculated the mean AFDM per specimen. This procedure was repeated 2–5× for each species, depending on how many specimens were available for incineration. Global mean AFDM ± SE for each species was then determined from these data.
Resultant AFDM values are in Table
The 63 species of caddisfly adults for which ash-free dry mass (AFDM) (± SE) was determined. Key: Year, year collected. #, number of specimens tested per incineration. N, the number of incinerations per species. M/F, whether male or female specimens were measured.
Taxon | Site | Year | # | N | M/F | AFDM (mg) | ± SE |
---|---|---|---|---|---|---|---|
APATANIIDAE | |||||||
Apatania zonella (Zetterstedt, 1840) | MI: Lk. Superior, 46.9083, -87.9225 | 2019 | 50 | 3 | M | 0.628 | 0.149 |
BRACHYCENTRIDAE | |||||||
Brachycentrus americanus (Banks, 1899) | MI: Fairbanks Cr., 44.0481, -85.6586 | 2014 | 50 | 3 | M | 0.745 | 0.104 |
Micrasema wataga Ross, 1938 | MN: Straight R., 46.8745, -95.0586 | 2000 | 300 | 3 | F | 0.094 | 0.026 |
DIPSEUDOPSIDAE | |||||||
Phylocentropus placidus (Banks, 1905) | MI: Nunn’s Cr., 46.0572, -84.5639 | 2010 | 35 | 3 | M | 0.418 | 0.064 |
GLOSSOSOMATIDAE | |||||||
Glossosoma nigrior Banks, 1911 | MI: Fairbanks Cr., 44.0481, -85.6586 | 2011 | 100 | 3 | F | 0.284 | 0.140 |
Protoptila maculata (Hagen, 1861) | MI: Manistee R., 44.2836, -85.8614 | 2010 | 500 | 3 | F | 0.030 | 0.008 |
GOERIDAE | |||||||
Goera stylata Ross, 1938 | MI: Fairbanks Cr., 44.0481, -85.6586 | 2011 | 100 | 6 | M | 0.495 | 0.074 |
HELICOPSYCHIDAE | |||||||
Helicopsyche borealis (Hagen, 1861) | MI: Black R., 45.1664, -84.3264 | 2015 | 250 | 6 | M | 0.223 | 0.042 |
HYDROPSYCHIDAE | |||||||
Cheumatopsyche campyla Ross, 1938 | MI: Tittabawasee R., 43.4811, -84.0931 | 2011 | 150 | 6 | M | 0.346 | 0.062 |
C. speciosa (Banks, 1904) | MN: Pine R., 46.5717, -94.0281 | 2000 | 150 | 3 | F | 0.245 | 0.054 |
Diplectrona modesta Banks, 1908 | MI: Fairbanks Cr., 44.0481, -85.6586 | 2014 | 50 | 3 | M | 0.502 | 0.071 |
Hydropsyche betteni Ross, 1938 | MI: Fairbanks Cr., 44.0481, -85.6586 | 2014 | 75 | 6 | M | 0.685 | 0.123 |
H. morosa (Hagen, 1861) | MI: Au Sable R., 44.6599, -84.1292 | 2011 | 100 | 3 | M | 0.392 | 0.110 |
H. simulans Ross, 1938 | MN: Chippewa R., 45.9408, -95.7383 | 2000 | 50 | 3 | M | 0.712 | 0.104 |
H. sparna Ross, 1938 | MI: Mountain St., 46.8692, -87.8933 | 2019 | 100 | 3 | M | 0.452 | 0.099 |
Macrostemum zebratum (Hagen, 1861) | WI: Peshtigo R., 45.2325, -88.0136 | 2015 | 50 | 3 | M | 0.884 | 0.159 |
Parapsyche apicalis (Banks, 1908) | MI: Fairbanks Cr., 44.0481, -85.6586 | 2011 | 50 | 3 | M | 0.472 | 0.066 |
Potamyia flava (Hagen, 1861) | IN: Ohio R., 37.7783, -87.9468 | 2018 | 200 | 6 | M | 0.399 | 0.072 |
HYDROPTILIDAE | |||||||
Agraylea multipunctata Curtis, 1834 | IN: Ohio R., 37.7783, -87.9468 | 2018 | 500 | 3 | F | 0.029 | 0.004 |
Hydroptila xera Ross, 1938 | MI: Two-hearted R., 46.6419, -85.4792 | 2011 | 500 | 3 | M | 0.017 | 0.003 |
Orthotrichia aegerfasciella (Chambers, 1873) | MI: Manistee R., 44.2836, -85.8614 | 2010 | 500 | 4 | M | 0.011 | 0.003 |
LEPIDOSTOMATIDAE | |||||||
Lepidostoma bryanti (Banks, 1908) | MI: Fairbanks Cr., 44.0481, -85.6586 | 2011 | 50 | 6 | M | 0.452 | 0.115 |
L. togatum (Hagen, 1861) | MI: Black R., 45.1664, -84.3264 | 2015 | 50 | 6 | M | 0.469 | 0.108 |
LEPTOCERIDAE | |||||||
Ceraclea arielles (Denning, 1942) | MI: Pine R., 44.1339, -85.6956 | 2010 | 150 | 3 | M | 0.318 | 0.054 |
C. resurgens (Walker, 1852) | MI: Mountain St., 46.8692, -87.8933 | 2019 | 75 | 3 | M | 0.712 | 0.459 |
C. tarsipunctata (Vorhies, 1909) | MI: Manistee R., 44.2836, -85.8614 | 2010 | 100 | 6 | M | 0.681 | 0.409 |
C. transversa (Hagen, 1861) | MN: North Brule R., 48.0076, -90.4169 | 2001 | 100 | 3 | M | 0.695 | 0.140 |
Leptocerus americanus (Banks, 1899) | MI: Saint Joseph R., 41.8361, -84.4772 | 2015 | 100 | 6 | M | 0.235 | 0.035 |
Mystacides interjecta (Banks, 1914) | MI: Benton Lk., 43.6718, -85.8916 | 2011 | 100 | 3 | M | 0.321 | 0.055 |
Nectopsyche candida (Hagen, 1861) | MI: Manistee R., 44.2836, -85.8614 | 2010 | 100 | 3 | M | 0.594 | 0.107 |
N. pavida (Hagen, 1861) | IN: Elkhart R., 41.5815, -85.8439 | 2018 | 100 | 3 | F | 0.254 | 0.116 |
Oecetis avara (Banks, 1895) | MI: Sturgeon R., 46.5689, -88.6564 | 2011 | 100 | 6 | M | 0.418 | 0.135 |
O. inconspicua (Walker, 1852) | MI: Bush Lk., 45.1919, -84.3177 | 2015 | 100 | 6 | M | 0.453 | 0.145 |
Setodes incertus (Walker, 1852) | MI: Big Sable R., 44.1176, -86.2010 | 2014 | 150 | 3 | M | 0.192 | 0.035 |
Triaenodes tardus Milne, 1934 | MN: Bush Lk., 45.1919, -84.3177 | 2015 | 50 | 3 | M | 0.595 | 0.166 |
LIMNEPHILIDAE | |||||||
Anabolia bimaculata (Walker, 1852) | MI: Silver Lk., 45.2042, -84.3117 | 2015 | 15 | 3 | M | 2.413 | 0.531 |
A. consocia (Walker, 1852) | MI: Fairbanks Cr., 44.0481, -85.6586 | 2011 | 15 | 3 | M | 1.849 | 0.407 |
Hydatophylax argus (Harris, 1869) | MI: Fairbanks Cr., 44.0481, -85.6586 | 2010 | 5 | 6 | F | 6.521 | 1.655 |
Limnephilus indivisus Walker, 1852 | MI: Fairbanks Cr., 44.0481, -85.6586 | 2012 | 10 | 3 | M | 2.295 | 0.487 |
Nemotaulis hostilis (Hagen, 1873) | MI: Fairbanks Cr., 44.0481, -85.6586 | 2012 | 7 | 3 | F | 5.515 | 0.827 |
Onocosmoecus unicolor (Banks, 1897) | MI: Salmon Trout R., 46.8485, -87.7989 | 2019 | 25 | 3 | M | 2.357 | 0.604 |
Platycentropus radiatus (Say, 1824) | MI: Fairbanks Cr., 44.0481, -85.6586 | 2013 | 8 | 3 | M | 3.973 | 0.596 |
Pycnopsyche antica (Walker, 1852) | MI: Fairbanks Cr., 44.0481, -85.6586 | 2013 | 25 | 6 | M | 2.263 | 0.354 |
P. guttifera (Walker, 1852) | MI: Fairbanks Cr., 44.0481, -85.6586 | 2012 | 25 | 6 | M | 2.199 | 0.396 |
P. lepida (Hagen, 1861) | MI: Mountain St., 46.8692, -87.8933 | 2019 | 25 | 3 | M | 2.095 | 0.342 |
MOLANNIDAE | |||||||
Molanna blenda Sibley, 1926 | MI: Fairbanks Cr., 44.0481, -85.6586 | 2011 | 50 | 3 | M | 0.686 | 0.099 |
M. uniophila Vorhies, 1909 | MI: Howe Lk. 46.8932, -87.9436 | 2019 | 50 | 3 | M | 0.715 | 0.122 |
ODONTOCERIDAE | |||||||
Psilotreta indecisa (Walker, 1852) | MI: Mountain St., 46.8692, -87.8933 | 2019 | 50 | 3 | M | 0.702 | 0.179 |
PHILOPOTAMIDAE | |||||||
Chimarra obscurra (Walker, 1852) | MI: Livermore Cr., 42.4457, -84.0420 | 2009 | 200 | 6 | M | 0.354 | 0.026 |
C. socia (Hagen, 1861) | MI: Sturgeon R., 46.5689, -88.6564 | 2011 | 200 | 6 | M | 0.402 | 0.082 |
Dolophilodes distinctus (Walker, 1852) | MI: Fairbanks Cr., 44.0481, -85.6586 | 2013 | 100 | 3 | M | 0.394 | 0.067 |
PHRYGANEIDAE | |||||||
Agrypnia improba (Hagen, 1873) | MI: Goose Pond, 45.7434, -84.8975 | 2011 | 7 | 3 | M | 3.059 | 0.551 |
Banksiola crotchi Banks, 1844 | MI: Fairbanks Cr., 44.0481, -85.6586 | 2013 | 25 | 6 | M | 1.371 | 0.412 |
Phryganea cinerea Walker, 1852 | MI: Fairbanks Cr., 44.0481, -85.6586 | 2011 | 5 | 6 | M | 6.846 | 1.504 |
Ptilostomis ocellifera (Walker, 1852) | MI: Fairbanks Cr., 44.0481, -85.6586 | 2011 | 5 | 6 | M | 7.169 | 1.367 |
P. semifasciata (Say, 1828) | MI: Slapneck Cr., 46.3331, -86.9369 | 2011 | 5 | 6 | M | 7.217 | 2.073 |
POLYCENTROPODIDAE | |||||||
Holocentropus interruptus Banks, 1914 | MI: Rockwell Lk., 44.0445, -85.6476 | 2011 | 50 | 3 | M | 0.511 | 0.982 |
Nyctiophylax affinis (Banks, 1897) | MI: Benton Lk., 43.6718, -85.8916 | 2011 | 200 | 3 | M | 0.105 | 0.018 |
Polycentropus pentus Ross, 1941 | MI: Fairbanks Cr., 44.0481, -85.6586 | 2014 | 75 | 3 | M | 0.418 | 0.092 |
PSYCHOMYIIDAE | |||||||
Psychomyia flavida Hagen, 1861 | WI: Red R., 44.8022, -88.6711 | 2015 | 500 | 6 | F | 0.038 | 0.005 |
RHYACOPHILIDAE | |||||||
Rhyacophila fuscula (Walker, 1852) | MI: Miners R., 46.4747, -86.5314 | 2011 | 75 | 6 | M | 1.402 | 0.355 |
SERICOSTOMATIDAE | |||||||
Agarodes distinctus (Ulmer, 1905) | MI: Howe Lk. 46.8932, -87.9436 | 2019 | 25 | 6 | M | 0.795 | 0.127 |
Thremmatidae | |||||||
Neophylax concinnus MacLachlan, 1871 | MI: Miners R., 46.4747, -86.5314 | 2019 | 75 | 3 | M | 0.329 | 0.071 |
The lack of previous research on the AFDM weights of adult caddisflies renders direct comparisons to other results impossible. Even indirect comparisons are difficult. Of the caddisflies previously weighed via dry mass calculation, none are of the same species that we weighed. Four species: Agrypnia deflata (Milne) (
Some weight differences between our specimens and those of other studies may also be due to actual variation between specimens. Several studies have reported 2–5× differences in dry mass between conspecific specimens in the same study due to differences in environmental conditions, larval food quality, or emergence timing (
This increased precision of using AFDM instead of specimen counting in ecological calculations can be observed when analyzing a previously published data set (
Comparison of mean specimen abundance (% of total specimens) and AFDM biomass (% of total biomass) for the caddisfly FFGs of a Michigan first order stream, based on 13 blacklight samples from each of 5 sites collected weekly from June to August 2012 (
These data allow, for the first time, the use of biomass data when assessing stream conditions using adult Trichoptera. Further research will be needed on intra- and inter-population biomass variation within a region. Further, the weights of the fairly high-latitude populations measured in our region may be different than lower latitude populations of the same species. It is our hope that similar studies are conducted in other areas of the US and elsewhere to further increase the value of the adult caddisflies as a biological monitoring taxon.
Collection of specimens utilized in this study was supported by a US Environmental Protection Agency Science to Achieve Results Fellowship, grants from the Minnesota Nongame Research Program and the Huron Mountains Wildlife Foundation, and funds from the Hillsdale College biology department. We thank Tony Swinehart for use of his laboratory and equipment. We thank Ralph Holzenthal and the University of Minnesota Insect Museum for donating specimens of several species. The valuable comments of Desi Robertson and Andy Rasmussen improved earlier versions of the manuscript. This is paper #22 of the GH Gordon Biological Station Research Series.