ArticlePDF Available

Length-dry mass relationships for a typical shredder in Brazilian streams (Trichoptera: Calamoceratidae)

Authors:

Abstract and Figures

The aims of this study were to determine which linear body dimensions are best suitable and which mathematical functions can be used to describe length–dry mass relationships for a population of Phylloicus sp. (Trichoptera: Calamoceratidae) larvae. We measured three linear body dimensions (body length, head capsule width and interocular distance) of 54 larvae to use as dry mass predictors. For the description of length–dry mass relationships we used linear, exponential and power function models. Body length provided the best fitted equations to estimate biomass, followed by head capsule width and interocular distance. The highest coefficients of determination were found in power function and exponential models. These relationships can be useful to determine the growth rate and/or secondary production of Phylloicus larvae in future laboratory experiments, as well as to understand the importance of these shredders in the energy flux of shaded tropical streams.
Content may be subject to copyright.
Length–dry mass relationships for a typical shredder in Brazilian
streams (Trichoptera: Calamoceratidae)
Ba
´rbara Becker, Marcelo S. Moretti and Marcos Callisto*
Universidade Federal de Minas Gerais, Instituto de Cie
ˆncias Biolo
´gicas, Departamento de
Biologia Geral, Laborato
´rio de Ecologia de Bentos, Belo Horizonte, MG, Brasil
(Received 29 June 2008; final version received 15 December 2008)
The aims of this study were to determine which linear body dimensions are best
suitable and which mathematical functions can be used to describe length–dry
mass relationships for a population of Phylloicus sp. (Trichoptera: Calamocer-
atidae) larvae. We measured three linear body dimensions (body length, head
capsule width and interocular distance) of 54 larvae to use as dry mass predictors.
For the description of length–dry mass relationships we used linear, exponential
and power function models. Body length provided the best fitted equations to
estimate biomass, followed by head capsule width and interocular distance. The
highest coecients of determination were found in power function and
exponential models. These relationships can be useful to determine the growth
rate and/or secondary production of Phylloicus larvae in future laboratory
experiments, as well as to understand the importance of these shredders in the
energy flux of shaded tropical streams.
Keywords: size-mass equations; biomass estimation; linear body dimensions;
Phylloicus; tropical shredders
Introduction
Biomass of aquatic macroinvertebrates is important to determine growth rates and/
or secondary production, as well as to understand life histories, seasonal patterns
and trophic relationships between functional feeding groups (Benke 1996; Burgherr
and Meyer 1997). Data on macroinvertebrate biomass can also be useful in
colonisation studies or quantifying the role of detritivores on leaf decomposition
(Cressa 1999).
Among the dierent approaches to biomass determination, the most common is
the direct weighing of individual specimens (Dermott and Paterson 1974; Smock
1980; Meyer 1989). However, this approach is often very time consuming, and prone
to error if the insects have been previously stored in chemical preservatives (e.g.
formalin or alcohol), which can cause alterations in their dry mass (Donald and
Paterson 1977; Downing and Rigler 1984; Kato and Miyasaka 2007). Direct
determination of dry mass has the added disadvantage of rendering the specimen
*Corresponding author. Email: callisto@icb.ufmg.br
Aquatic Insects
Vol. 31, No. 3, September 2009, 227–234
ISSN 0165-0424 print/ISSN 1744-4152 online
!2009 Taylor & Francis
DOI: 10.1080/01650420902787549
http://www.informaworld.com
Downloaded By: [Callisto, Marcos] At: 14:37 7 August 2009
useless for further examination as a result of the drying process (Towers, Henderson
and Veltman 1994).
An alternative to avoid such disadvantages is to estimate the biomass indirectly,
using length–dry mass conversions (Gould 1966; Peters 1983; Burgherr and Meyer
1997; Benke, Huryn, Smock and Wallace 1999). Estimating dry mass indirectly from
linear body dimensions (e.g. body length, head capsule width) is more rapid than
direct mass determination, particularly for small invertebrates. Moreover, in
laboratory experiments assessing invertebrate feeding behaviour, this approach allows
the estimation of initial biomass without stressing and/or killing the organisms.
Length–dry mass relationships have been used to estimate the biomass of
invertebrates from dierent geographical locations and of taxa with similar body
shapes (Johnston and Cunjak 1999). Most of the length–dry mass relationships for
stream invertebrates were estimated for North American and European taxa (Smock
1980; Meyer 1989) and, until now, only a few data were proposed for the tropical
region. Furthermore, previous studies suggested the need to use taxa-specific
relationships because they are more precise, once dierent taxa may dier in body
shape and volume (Schoener 1980; Smock 1980; Gowing and Recher 1985; Cressa
1986).
Only few invertebrate taxa have been mentioned as shredders in neotropical
streams. Among them, larvae of the genus Phylloicus Mu
¨ller, 1880 (Trichoptera:
Calamoceratidae) are well distributed throughout Latin America and, in some
streams, can be found easily on leaf patches with low water current (Prather 2003).
Because these larvae are also easy to manipulate and keep alive in laboratory
conditions, they have been used in many experiments (e.g. feeding preference,
growth, survival and case building) that aimed to better understand the behaviour of
shredders and their influence on leaf decomposition in tropical streams (Grac¸ a et al.
2001; Rinco
´n and Martı
´nez 2006).
In this study, we analysed the length–dry mass relationships for a population
of Phylloicus sp. by using three dierent regression functions (linear, power
and exponential) and three body dimensions in order to determine the best
relationship.
Materials and methods
Phylloicus sp. larvae were collected on July 2007 in Taboo
˜es spring (2080303800S–
4480300300W), located in the Serra do Rola Moc¸ a State Park, Minas Gerais State,
southeastern Brazil. The Taboo
˜es spring is inside a forest fragment, presenting a well
developed riparian area, which forms a closed canopy. Leaves fall throughout the
year and accumulate in the streambed.
Larvae were found visually, collected with a hand net, and taken to the
laboratory in an isothermic box with stream water. In the laboratory, undamaged
individuals of the same morphospecies were carefully removed from their cases and
placed individually in Petri dishes. Three linear body dimensions were chosen among
the most common used as biomass predictors: body length, head capsule width and
interocular distance (Meyer 1989). Body length (BL) was measured as the distance
from the anterior of the head to the posterior of the last abdominal segment. Head
capsule width (HW) was measured across the widest section of the head. Interocular
distance (ID) was measured as the minimum distance between eyes, parallel to head
width. Body dimensions were measured to the nearest 0.1 mm with a Zeiss dissecting
228 B. Becker et al.
Downloaded By: [Callisto, Marcos] At: 14:37 7 August 2009
microscope fitted with an ocular micrometer (magnification: 8 x for BL measure-
ments and 50 x for HW and ID measurements). Animals were then placed
individually in pre-weighed aluminium foils, dried at 608C for 48 h (Meyer 1989), left
to cool in a desiccator, and their dry mass (DM) was measured to the nearest 0.1 mg.
Three regression models were calculated for the three Phylloicus body
dimensions, using the method of least squares. The fit of regression equations was
judged by the coecient of determination (r
2
), the significance level (p, obtained
from regression ANOVA) and residual analysis. All statistical analyses were
performed based on Zar (1999).
Results
Body dimensions measures and dry weights of 54 larvae were used for statistical
analyses. Phylloicus dry mass presented the highest coecient of variation, with
values ranging from 1.3 to 26.6 mg (Table 1). Among body dimensions, body length
presented higher range (10.4–28.9 mm) and coecient of variation (Table 1).
The following regression models were chosen because they provided the best fits.
Conversion of Phylloicus body dimensions to dry mass was determined by linear (1),
exponential (2) and power function (3) models or its logarithmic equivalents:
DM ¼aþb#Lð1Þ
DM ¼a#ebL in linear form:ln DM ¼ln aþb#Lð Þ ð2Þ
DM ¼a#Lbin linear form:ln DM ¼ln aþb#ln Lð Þ ð3Þ
where a/bare regression constants, DM is dry mass, Lis the linear body dimension
(BL, HW, ID) and eis a mathematical constant (Euler’s number: 2.718).
The parameters of Equations (1), (2) and (3) are listed in Table 2. All
body dimensions showed a very high level of significance in the three models
(p50.01). Body length provided the best relationships to estimate biomass
(Table 2), followed by head capsule width and interocular distance. These
relationships were best fitted by power function and exponential models that
presented very similar coecients of determination to each body dimension. Figure 1
shows the relations of dry mass as a function of body length, head capsule width and
interocular distance for Phylloicus larvae. The regression lines and curves were given
by power function.
Table 1. Ranges, mean, standard deviation (SD) and coecient of variation (CV, in
percentage) for body length, head capsule width, interocular distance (mm) and dry mass (mg)
of Phylloicus sp. larvae; n¼54. CV ¼(SD/mean)6100.
Range Mean SD CV
Body length 10.4–28.9 16.7 2.7 16.2
Head capsule width 0.8–1.5 1.3 0.2 14.0
Interocular distance 0.6–1.1 1.0 0.1 14.3
Dry mass 1.3–26.6 12.1 6.7 54.2
Aquatic Insects 229
Downloaded By: [Callisto, Marcos] At: 14:37 7 August 2009
Discussion
Even though all relationships between body dimensions and biomass were highly
significant, body length was the best predictor, explaining 75–76% of the variation in
mass. This linear body dimension is widely used for determining length–dry mass
relationships of aquatic invertebrates (e.g. Smock 1980; Towers et al. 1994; Burgherr
and Meyer 1997) mainly because it has a broader measuring range. Body length also
provides slightly higher coecients of determination than head capsule width and
interocular distance (Gonza
´lez, Basaguren and Pozo 2002).
Although body length usually gives the best relationships, some authors (see
Cressa 1999; Marchant and Hehir 1999; Gonza
´lez et al. 2002) prefer to use other
linear body dimensions, like head capsule width, case width, pronotum length or
tarsus length. This probably owes to the fact that, among other reasons, these
structures are sclerotised and less subject to distortion or breakage under
manipulation than body length. In addition, Becker (2005) found that pronotum
length is the best measurement to distinct larval instars of Agapetus fuscipes
(Trichoptera) in a German first-order stream. In the present study, larvae were
measured on the same day they had been sampled. So, all measurements were done
on fresh, undamaged and completely stretched animals, which allowed a precise and
reliable determination of the three studied body dimensions.
The exponential and power function models did not dier between the body
dimensions determined. Most authors found the highest fit between body length and
dry mass when they use the power function model (e.g. Smock 1980; Meyer 1989;
Burgherr and Meyer 1997) but exponential regressions have also been used by
Dudgeon (1995) and Pera
´n, Velasco and Milla
´n (1999) for length–dry mass
relationships of Hydrocyphon (Coleoptera) and Caenis luctuosa (Ephemeroptera),
respectively. Wenzel, Meyer and Schwoerbel (1990) pointed out that dierences
between the results obtained using dierent regression models are low and they
decrease when a higher number of animals is used. Although power function is more
often used, the exponential model should not be discarded when looking for the best
fit of length–dry mass relationships.
Table 2. Parameters (with 95% confidence intervals) of the linear, exponential and power
function models for the relationship between a linear body dimension (L¼body length [BL],
head capsule width [HW] or interocular distance [ID], in mm) and dry mass (DM, in mg) of
Phylloicus sp. larvae.
Function Conversion a ln a b r
2
Linear BL.!DM 720.24 +3.44 1.93 +0.20 0.64**
DM ¼aþb#L HW.!DM 721.20 +4.58 26.49 +3.61 0.51**
ID.!DM 719.97 +4.62 33.51 +4.78 0.49**
Exponential BL.!DM 71.63 +0.32 0.23 +0.02 0.75*
ln DM ¼
ln a þb#L
HW.!DM 72.05 +0.40 3.45 +0.32 0.70**
ID.!DM 72.00 +0.39 4.48 +0.41 0.70**
Power BL.!DM 77.73 +0.77 3.58 +0.28 0.76*
ln DM ¼
ln a þb#ln L
HW.!DM 1.43 +0.09 3.95 +0.35 0.71**
ID.!DM 2.50 +0.06 3.84 +0.34 0.71**
a, b ¼regression constants, r
2
¼coecient of determination (*p50.01, **p50.001). n¼54.
230 B. Becker et al.
Downloaded By: [Callisto, Marcos] At: 14:37 7 August 2009
In practice, when interpreting a length–dry mass regression equation, ‘‘b’’
values represent the rate of increase (i.e. slope) of dry weight against length in a
linear relationship, whereas the constant ‘‘a’’ only represents the dry mass of
an organism at a unit length (i.e. 1 mm). It is known that for tropical aquatic
insects the constant bfalls short of the expected value of 3, which means that
body mass of insects is more influenced by surface than by volume (Engelmann
1961). Our results support those from Cressa (1999) who found that Phylloicus
sp. is one of the few taxa of tropical invertebrates whose slope is higher than 3,
so it is possible that in this genus volume could influence body mass more than
surface.
Figure 1. Scatter diagrams of (A) dry mass versus body length, (B) head capsule width and
(C) interocular distance on normal coordinates (¤) as well as on logarithmic coordinates (.)
for Phylloicus sp. larvae. The regression equations (power function) are DM ¼a#L
b
and ln
DM ¼ln aþb#ln L.
Aquatic Insects 231
Downloaded By: [Callisto, Marcos] At: 14:37 7 August 2009
Some variations in length–dry mass relationships for the populations of the
same species, but from dierent locations, can be caused by physical–chemical
dierences of the environment, trophic conditions or genetics. In this way, it is
recommended to determine the relationships for populations under study or use
relationships that were determined for populations from the same streams and/or
regions. For example, Rinco
´n and Martinez (2006), studying the growth rates of
Phylloicus in laboratory experiments, used the empirical relationship described by
Cressa (1999) who had studied populations from a similar region of Venezuela.
On the other hand, length–dry mass relationships are not much aected by
seasons, as shown by Kato and Miyasaka (2007). These authors suggested that it
is not necessary to measure larvae in dry and wet seasons to have a consistent
relationship.
When sampling organisms to determine length–dry mass relationships, one
must be sure that organisms from dierent sizes (cohorts) have been collected. If
not, only part of the logistic curve of population growth is quantified and the
resulting relationships may not represent the whole population (Begon, Mortimer
and Thompson 1996). In this study, if we consider Dyar’s law, an empirical law
that suggests an increase of 1.5 in growth at each instar (Wigglesworth 1972), and
the ranges of each body dimension measured, we can infer that only larvae from
the last two instars were sampled. Based on this, our equations were determined
with data from the right side of the curvilinear relationships between dry mass and
body dimensions of this population of Phylloicus (see Majecki, Grzybkowska and
Reddy 1997). On the other hand, as we have been monitoring this population for
several months, larvae used in this study presented the same range of size of the
ones that are found visually in most part of the year, suggesting that our equations
were adequate to determine the dry mass of larvae destined to laboratory
experiments.
In conclusion, the length–dry mass relationships presented here can be useful to
determine the growth rate and/or secondary production of Phylloicus. Besides, our
results also reinforce the necessity of more studies focusing on the life cycles of
aquatic insects in the tropical region. We do hope that the present study encourages
future research assessing the population dynamics of tropical shredders, as well as to
understand the importance of these individuals on leaf processing, trophic
relationships, colonisation rates, and even to compare populations within and
between habitats.
Acknowledgements
This study was supported by FAPEMIG, IEF-MG, CoPASA, CNPq, CAPES, Eawag, US Fish
and Wildlife Service. We appreciated the help of our laboratory colleagues Lurdemar Tavares
and Juliana Franc¸ a during field and laboratory activities. We are also thankful to Joa
˜o Jose
´
Leal, Leandro Oliveira, Vicenc¸ Acun
˜a and two anonymous reviewers who provided useful
comments on the manuscript.
References
Becker, G. (2005), ‘Life cycle of Agapetus fuscipes (Trichoptera, Glossosomatidae) in a first-
order upland stream in central Germany’, Limnologica, 35, 52–60.
Begon, M., Mortimer, M., and Thompson, D.J. (1996), Population Ecology: A Unified Study
of Animals and Plants (3rd ed.), Oxford: Blackwell Science.
Benke, A. (1996), ‘Secondary production of macroinvertebrates’, in Methods in Stream
Ecology, eds. F.R. Hauer and G.A. Lamberti, Academic Press: New York, pp. 557–578.
232 B. Becker et al.
Downloaded By: [Callisto, Marcos] At: 14:37 7 August 2009
Benke, A., Huryn, A., Smock, L., and Wallace, J. (1999), ‘Length–mass relationships for
freshwater macroinvertebrates in North America with particular reference to the
southeastern United States’, Journal of the North American Benthological Society, 18,
308–343.
Burgherr, P., and Meyer, E.I. (1997), ‘Regression analysis of linear body dimensions vs. dry
mass in stream macroinvertebrates’, Archiv fu
¨r Hydrobiologie, 139, 101–112.
Cressa, C. (1986), ‘Estimaciones de peso seco en funcio
´n de la longitud cefa
´lica y clases de
taman
˜o en Campsurus sp. (Ephemeroptera, Polymitarcidae)’, Acta Cientı´fica Venezolana,
37, 170–173.
Cressa, C. (1999), ‘Dry mass estimates of some tropical aquatic insects’, Revista de Biologı´a
Tropical, 47, 133–141.
Dermott, R.M., and Paterson, C.G. (1974), ‘Determining dry weight and
percentage dry matter of chironomid larvae’, Canadian Journal of Zoology, 52,
1243–1250.
Donald, G.L., and Paterson, C.G. (1977), ‘Eects of preservation on wet weight biomass of
chironomid larvae’, Hydrobiologia, 53, 75–80.
Downing, J.A., and Rigler, F.H. (1984), A Manual on Methods for the Assessment of
Secondary Productivity in Fresh Waters, Blackwell: Oxford.
Dudgeon, D. (1995), ‘Life history, secondary production and microdistribution of
Hydrocyphon (Coleoptera: Scirtidae) in a tropical forest stream’, Archiv fu
¨r Hydrobiologie,
133, 261–271.
Engelmann, M.D. (1961), ‘The role of soil arthropods in the energetics of an old field
community’, Ecological Monographs, 31, 221–238.
Gonza
´lez, J.M., Basaguren, A., and Pozo, J. (2002), ‘Size–mass relationships of stream
invertebrates in a northern Spain stream’, Hydrobiologia, 489, 131–137.
Gould, S. (1966), ‘Allometry and size in ontogeny and phylogeny’, Biological Research, 41,
587–640.
Gowing, G., and Recher, H.F. (1985), ‘Length-weight relationships for
invertebrates from forest in south-eastern New South Wales’, Australian Journal of
Ecology, 9, 5–8.
Grac¸ a, M.A.S., Cressa, C., Gessner, M.O., Feio, M.J., Callies, K.A., and Barrios, C. (2001),
‘Food quality, feeding preferences, survival and growth of shredders from temperate and
tropical streams’, Freshwater Biology, 46, 947–957.
Johnston, T., and Cunjak, R. (1999), ‘Dry mass-length relationships for benthic insects: a
review with new data from Catamaran Brook, New Brunswick, Canada’, Freshwater
Biology, 41, 653–674.
Kato, M.G., and Miyasaka, H. (2007), ‘Length–weight relationships of four predatory
stonefly species in Japan’, Limnology, 8, 171–174.
Majecki, J., Grzybkowska, M., and Reddy, R. (1997), ‘Density, production and life cycle of
Brachycentrus subnubilus Curtis (Trichoptera: Brachycentridae) in a lowland river, Central
Poland’, Hydrobiologia, 354, 51–56.
Marchant, R., and Hehir, G. (1999), ‘Growth, production and mortality of two species of
Agapetus (Trichoptera: Glossosomatide) in the Acheron River, south-east Australia’,
Freshwater Biology, 42, 655–671.
Meyer, E. (1989), ‘The relationship between body length parameters and dry mass in running
water invertebrates’, Archiv fu
¨r Hydrobiologie, 117, 191–203.
Pera
´n, A., Velasco, J., and Milla
´n, A. (1999), ‘Life cycle and secondary production of Caenis
luctuosa (Ephemeroptera) in a semiarid stream (Southeast Spain)’, Hydrobiologia, 400,
187–194.
Peters, R.H. (1983), The Ecological Implications of Body Size, Cambridge: Cambridge
University Press.
Prather, A.L. (2003), ‘Revision of the Neotropical caddisfly genus Phylloicus (Trichoptera:
Calamoceratidae)’, Zootaxa, 275, 1–214.
Rinco
´n, J., and Martı
´nez, I. (2006), ‘Food quality and feeding preferences of Phylloicus sp.
(Trichoptera: Calamoceratidae)’, Journal of the North American Benthological Society, 25,
209–215.
Schoener, T.W. (1980), ‘Length–weight regressions in tropical and temperate forest-
understorey insects’, Annals of the Entomological Society of America, 73, 106–109.
Aquatic Insects 233
Downloaded By: [Callisto, Marcos] At: 14:37 7 August 2009
Smock, L.A. (1980), ‘Relationships between body size and biomass of aquatic insects’,
Freshwater Biology, 10, 375–383.
Towers, D.J., Henderson, I.M., and Veltman, C.J. (1994), ‘Predicting dry weight of New
Zealand aquatic macroinvertebrates from linear dimensions’, New Zealand Journal of
Marine and Freshwater Research, 28, 159–166.
Wenzel, F., Meyer, E., and Schwoerbel, J. (1990), ‘Morphometry and biomass determination
of dominant mayfly larvae (Ephemeroptera) in running waters’, Archiv fu
¨r Hydrobiologie,
118, 31–46.
Wigglesworth, V.B. (1972), The Principles of Insect Physiology, London: Chapman & Hall.
Zar, J.H. (1999), Biostatistical Analysis, Englewood Clis, NJ: Prentice Hall.
234 B. Becker et al.
Downloaded By: [Callisto, Marcos] At: 14:37 7 August 2009
... 35, e5 Mährlein et al., 2016). Furthermore, length-body mass relationships should be taxon-specific, as there may be a different relationship for each taxon (Baumgärtner & Rothhaupt, 2003;Becker et al., 2009). Therefore, studies on the biomass of aquatic insects should consider the specificity of each region. ...
... The measured individuals were placed individually in pre-weighed porcelain crucibles, dried in an oven at 60°C for 48 h (Becker et al., 2009), allowed to cool in a desiccator and their dry mass measured on a balance with ± 0.001 g accuracy. Subsequently, to estimate the ash weight, the individuals were incinerated in a muffle furnace at 550°C for 4 hours, and their ash mass was measured by the same procedure. ...
... Our results showed that the power models presented a high correlation coefficient, explaining 78% to 90% of the variation in biomass of Leptonema larvae as a function of the allometric measurements used (body length, interocular distance, horizontal head size and vertical head size). In fact, our results are in line with other studies in the tropical region (e.g., Becker et al., 2009;Silva et al., 2010). This reinforces that power models for length-mass of aquatic macroinvertebrates provide satisfactory results for the relationship between body dimensions and biomass of freshwater invertebrates, including Leptonema larvae. ...
Article
Full-text available
Aim: The objectives of this study were to evaluate which allometric measurements of Leptonema larvae are most suitable in order to develop mathematical equations to describe biomass relationships for the population of this taxon in a reference condition headwater stream. Methods: We measured four body dimensions (body length, interocular distance, horizontal width of cephalic capsule and vertical width of the cephalic capsule) of 65 Leptonema larvae, which were collected in February 2022, in the Taboões spring, Serra do Rola Moça State Park, Minas Gerais, using a Surber sampler. For the determination of allometric measurements, each individual was photographed under a dissecting stereomicroscope (Leica M80) equipped with a digital camera. Each photographed specimen's length was measured using the Motic Image Plus 2.0 software. After measuring the linear body dimension and direct measurement of the biomass, we used these values to calculate the length-mass mathematical equations. To the equations use power models: DM = a Lb, where a/b are constants, DM is the dry mass, L is the linear body dimension. Results: Among body dimensions of Leptonema larvae, body length showed the greatest range of variation, with values ranging from 4.03 to 25.28 mm, followed by head capsule vertical width (0.51-2.69 mm), head capsule horizontal width (0.55-2.22 mm) and interocular distance (0.24-1.88 mm). Our results show that body length provided the best-fitting equation for estimating biomass (R 2 = 0.90). However, we observed a close fit between the other allometric measures, including high coefficients of determination, head capsule horizontal width (R 2 = 0.85), interocular distance (R 2 = 0.82), head capsule vertical width (R 2 = 0.78). Conclusions: These results will be useful in providing the best allometric measurement and equations to estimate the biomass of Leptonema larvae from the tropics.
... Most previous studies relied on one a priori chosen linear trait and rarely compared multiple traits in length-mass relationships (but see Wenzel et al. 1990, Becker et al. 2009, Shahbaz-Gahroee et al. 2021. Here, we compared 3 commonly used traits: head width, body length, and body width (= thorax width in mosquito larvae). ...
... These traits are not always equally suitable (Wenzel et al. 1990): the trait should be clearly delimited, correlate tightly with body mass, and preferably be planar to limit the measurement error when converting from a 3-dimensional structure to a 2-dimensional image (Cardini 2014). However, despite different accuracy in estimating the mass in lengthmass relationships (e.g., Becker et al. 2009), studies that deal with a comparison of morphological traits to each other are quite rare, and thus, our study brings additional insight into this field. ...
Article
Full-text available
Body mass underpins many ecological processes at the level of individuals, populations, and communities. Often estimated in arthropods from linear morphological traits such as body length or head width, these relationships can vary even between closely related taxa. Length–mass relationships of mosquito (Diptera: Culicidae) larvae are poorly known despite the importance of this family to disease and aquatic ecology. To fill this gap, we measured ontogenetic changes in linear traits (body length, head width, and thorax width) and dry and wet masses and estimated length- and width–mass relationships in larvae of 3 culicid species inhabiting different niches: the tropical Aedes albopictus (Skuse, 1894), the temperate Culex pipiens (Linnaeus, 1758), and the snowmelt Ochlerotatus punctor (Kirby, 1837). We compared our results with published length–mass allometries of other aquatic dipteran larvae. We showed that thorax width and body length, but not head width, reliably predicted body mass for our 3 species. The length–mass allometry slopes in aquatic dipterans varied considerably between and within families but were independent of phylogeny, specimen handling, preservation techniques, and data fitting methods. Slope estimates became less precise with decreasing sample size and size range. To obtain reliable estimates of the allometric slopes, we have thus recommended using data on all larval stages for intraspecific allometries and a wide range of species for interspecific allometries. We also cautioned against the indiscriminate use of length–mass allometries obtained for other taxa or collected at lower taxonomic resolutions, e.g., when using length–mass relationships to estimate biomass production at a given site.
... Invertebrates were removed from cases prior to the start of experiments to avoid the consumption of external sources of organic matter (e.g., consumption of case) (Rezende et al. 2015). Interocular distance (i.e., the minimum distance between the eyes (± 0.01 mm); for more details, see Becker et al. (2009) and Martins et al. (2014)) was measured for each larva at the beginning of the experiment to estimate increase or decrease on body size at the end of the experiment. Then, each larva was carefully placed in microcosms (15.5 × 15.5 × 12 cm, 2.883 cm 3 volume) with gravel substrate previously sterilized in an oven for 4 h at 550 °C (Rezende et al. 2021a) and filled with bottled water. ...
... where DMc is disc dry mass used for cased building by Phylloicus sp. at the end of the experiment (7 days). Finally, we quantified the intraocular distance of Phylloicus sp. by the difference between final and initial sizes (mm) divided by initial size (mm) (Meyer 1989;Becker et al. 2009;Martins et al. 2014). ...
Article
Changes in riparian vegetation can alter the input and quality of leaf litter in aquatic ecosystems, but the effects of these changes on litter fragmentation by invertebrate shredder communities in tropical streams remain poorly studied. The caddisfly genus Phylloicus Müller, 1880 (Trichoptera: Calamoceratidae) is highly abundant in Neotropical streams, representing a great part of shredder biomass, which uses the allochthonous litter as a food resource and for case-building. We investigated leaf consumption by Phylloicus sp. under different leaf conditioning (leached and unleached) and plant species (Eucalyptus grandis, Erythrina falcata, and Inga uruguensis). The effects of leaf conditioning and plant species were measured using microcosm treatments, with one free Phylloicus sp. larva per l microcosm, and a decomposition control to correct for microbial decomposition. Our study suggests that phosphorus and caloric values of leaf litter are more important than leaf hardness and nitrogen in driving leaf consumption by Phylloicus sp. On the one hand, higher consumption was observed in treatment with unleached leaves than in leached leaf treatment due to higher nutrient concentration and caloric values on unleached leaves. On the other hand, Phylloicus sp. larvae preferred leached leaves for case building over unleached leaves, as leached leaves are less prone to the activity of the decomposing community, thus lowering the need for constant case renewal. Finally, high litter consumption is not necessarily converted into biomass by Phylloicus sp. larvae. In this sense, Phylloicus sp. larvae showed selectivity for resources with high caloric content for consumption and low caloric content for case-building.
... The biomass of the larvae can be estimated indirectly, using different size parameters of the body, if a given parameter is related significantly to the body weight: body length, pronotum length, interocular distance, tibia length, and head capsule width (Gould 1966;Benke, Huryn, Smock, and Wallace 1999;Becker 2005;de Brito, Martins, Soares, and Hamada, 2015;Kiffer, Mendes, Rangel, Barbosa, Serpa, and Moretti 2016). In most studies, larval biomass was calculated from the relationship between body weight and body length or width of head capsule (Benke, Huryn, Smock, and Wallace 1999;Miyasaka et al. 2008;Becker, Moretti, and Callisto 2009). In our sample of larvae, we examined the width of head capsule as a measure of larval growth, which was expected to correlate with the biomass of the larvae. ...
Article
Apatania crymophila McLachlan, 1880 is the most abundant species among trichopterans of the Yenisei River. The density of its free-moving larvae associated with water moss in the Yenisei River downstream of the high-pressure Krasnoyarsk Hydroelectric Plant dam was studied from June through November 2022. The biomass of A. crymophila larvae best correlated with case length, and the correlation was well fitted by linear equation; case length was also used to distinguish instars in this study. The abundance and biomass of A. crymophila larvae on water moss increased most rapidly (0.12 and 0.18day−1, respectively) from early July to early August, when water temperature approached its seasonal maximum. The maximum value of the larval density on water moss was reached in October (1084 larvae per kg of fresh moss). The relationship between larval growth and water temperature suggests that altered environmental conditions in the Yenisei River downstream of the HEP can be considered as a factor favoring the abundancy of A. crymophila.
... (MedCalc Software Ltd., Belgium). These measurements served to estimate the dry weight of the individuals based on established relationships (Potter and Learner, 1974;Smock, 1980;Meyer, 1989;Benke et al., 1999;Pavlov and Zubina, 1990;Burgherr and Meyer, 1997;Johnston and Cunjak, 1999;Martín, 2001;Miserendino, 2001;Miyasaka et al., 2008;Becker et al., 2009). ...
Article
Full-text available
We compared the structural and functional attributes of Neotropical macroinvertebrate communities between two natural, two agricultural, and two urban streams. We quantified food resource availability, community structure, secondary production, and consumer-resource interaction strength, attributing them to environmental characteristics. In agricultural streams, secondary production was significantly reduced compared to natural streams, reaching the lowest reported estimates for tropical streams (0.08-0.69 g DW⋅m − 2 ⋅year − 1 in agricultural streams vs. 1.1-15.8 g DW⋅m − 2 ⋅year − 1 in natural streams). This decline was primarily attributed to habitat degradation caused by siltation and channelization, alongside stronger top-down pressure in agricultural streams. Conversely, urban streams exhibited a substantial increase in secondary production, ranking among the highest estimates for tropical streams (25.6-191.2 g DW⋅m − 2 ⋅year − 1). The high secondary production in urban streams was primarily due to substantial increases in fine benthic organic matter, likely originating from poorly treated wastewater. Along with higher resource availability, we also observed a lack of, or insufficient, top-down control by macroinvertebrate predators or fish, which may have promoted higher secondary production in urban streams. This study is the first to quantify land use impacts on energy flows in stream communities in a region particularly threatened by land use change. Future studies should address multifunctional responses to land use changes in both the tropical and temperate realms to shed new light on how stream ecosystem functions in different biomes respond to human impacts. Our findings highlight the urgent need for effective management strategies tailored to address the environmental characteristics of tropical stream ecosystems.
... In addition, the absence of these genera in the adult stage may be related to the habits of these organisms. For example, Phylloicus are usually active during the day, unlike most Trichoptera, which may explain their absence from nocturnal light traps (Becker et al. 2009;Ferreira et al. 2015). Therefore, although the number and richness of adult Trichoptera attracted by light traps were high, we are aware that our sampling device, like any other, is not free from biases and limitations. ...
Article
Biological communities have their biodiversity patterns affected by environmental, spatial, and biogeographic factors that vary from taxa to taxa, and often between life stages. This is especially true when there are differences in the habitat the species use in each of them. Individuals of the insect order Trichoptera are mostly aquatic in their larval stage and terrestrial in their adult stage, which may result in different behaviors and environmental requirements. Our goal was to evaluate the congruence between the larval and adult stages of Trichoptera in Amazonian streams regarding their abundance, richness, and assemblage composition. Additionally, we tried to identify the main environmental factors related to each life stage. For this, larvae and adults of Trichoptera were sampled in the same sites at 12 streams in the Caxiuanã National Forest, Pará state, Brazil. Adult assemblages had greater richness of genera and abundance of individuals than the larval ones, and there was no congruence in the genera composition between these life stages. Our results also showed that different environmental variables structured Trichoptera larvae and adults. Since the sampling of larvae and adults proved to be complementary in the studied streams, we advise that Trichoptera diversity surveys consider both life stages of these organisms.
... La estimación indirecta de la biomasa en peso seco a partir de la longitud total del cuerpo es mucho más rápida que la determinación directa por gravimetría, particularmente para pequeños invertebrados como los macroinvertebrados acuáticos. Por supuesto, este tipo de técnicas indirectas dan una aproximación al peso seco, pero permiten evaluar la biomasa sin someter a los organismos a estrés y sin sacrificar toda la comunidad (Becker et al. 2009). Las determinaciones directas requieren una gran cantidad de muestras o especímenes recolectados, con lo cual, si hay muchos muestreos, pueden llegar a ser destructivos y nocivos para las poblaciones. ...
Article
Full-text available
Existen dificultades para determinar el peso de los macroinvertebrados y conocer así su importancia energética dentro de un ecosistema acuático. Por lo tanto, el objetivo de este estudio fue hallar relaciones matemáticas entre la longitud y la biomasa en dos géneros de invertebrados, representativos en el humedal Jaboque (Engativá) Bogotá D. C., Colombia, que faciliten hallar el peso de los individuos. Para ello, se realizaron cuatro muestreos desde abril de 2009 a enero de 2010. Se obtuvieron ecuaciones para estimar el peso seco a partir de medidas de la longitud total del cuerpo, para los géneros Helobdella y Asellus. Las relaciones halladas entre longitud total y peso seco fueron significativas y explicaron al menos el 69% de la varianza, expresada en los coeficientes de correlación (r2 = 0,69 y 0,85). Estos modelos permitirán calcular la biomasa para determinar el crecimiento y la producción secundaria de estos taxones en posteriores estudios de laboratorio o de campo. También podrían ayudar a conocer la importancia de estos organismos en el flujo de energía en los ecosistemas acuáticos, principalmente en los humedales andinos urbanos, donde son muy abundantes.
Article
Mass or volume estimators using body surface areas can be useful for environmental quality biomonitoring programs. We propose models of biovolume estimate through biometric information of Aulophorus, Dero, Nais and Pristina. The samples were collected in 51 freshwater systems in the state of São Paulo, between 2004 and 2016. Specific dimensions for each genus of 1,814 total animals were measured and individual biovolume was calculated. The conversion model was performed by linear regression analysis between the biovolume and the specific dimensions, then transformed into a power function, established by allometric law. The biovolume of Aulophorus and Dero can be estimated by the diameter of segment four (DIV), by the equation Be = 0.775DIV4.387 (adjusted r2 = 0.84). For Nais species, the biovolume can be estimated by the diameter of segment seven (DVII), by the equation Be = 0.913DVII1.6 (adjusted r2 = 0.59). For Pristina, the biovolume can be estimate by the diameter of segment eight (DVIII), by the equation Be = 0.875DVIII2.11 (adjusted r2 = 0.69). The application of these models can help us better understand population structure, secondary production and flow of matter and energy in neotropical environments.
Article
Full-text available
We propose a field procedure for estimating the dry biomass of stream macroinvertebrates. Estimates are calculated using the mean values of the a and b regression coefficients from unpublished data and an extensive review of the relevant literature. The regression equation employed for calculating dry biomass is one that has been extensively used: Y = aXb, where Y = mg dry mass of an individual macroinvertebrate; X = mm total body length of an individual macroinvertebrate; a = intercept coefficient of the Y on X regression; and b = slope coefficient Y on X. The procedure was developed for use in the field, but dry mass estimates can also be made on preserved specimens. The case is made for presenting stream macroinvertebrate dry biomass data categorized by functional feeding groups (FFGs) and their component higher level taxa. The tables summarize the FFGs and their food resources, mean regression coefficients, dry biomass estimates for FFG-taxa by size and a comparison of their numerical-to-gravimetric surrogate FFG ratios to predict the stream environmental condition. A sizing template for rapidly sorting macroinvertebrates in the field is described. Thresholds for surrogate FFG ratios that directly predict measured stream ecosystem conditions are described.
Article
Full-text available
Antecedentes: El estudio de la morfometría de los insectos acuáticos es importante para comprender algunos procesos ecológicos como la ganancia de la biomasa, el tiempo de desarrollo de una especie y la dinámica de las cohortes. En Colombia, no existen estudios sobre relaciones talla-peso seco a nivel de especie en ninfas de Anacroneuria. Objetivos: El objetivo de este trabajo fue analizar algunas características morfológicas de ninfas de Anacroneuria marta y A. caraca para determinar la eventual existencia de funciones lineales que expliquen la relación entre algunas dimensiones del cuerpo (la longitud total y el ancho de la cabeza) versus el peso seco. Métodos: Los organismos se recolectaron en zonas de rápidos y pozos del río Gaira (Sierra Nevada de Santa Marta, Colombia) entre octubre de 2014 y marzo del 2015. Las ninfas se identificaron a nivel de especie y las entidades taxonómicas se confirmaron mediante un análisis discriminante utilizando mediciones de diez variables morfológicas de las ninfas inmaduras. En algunos organismos se obtuvo el peso seco, el cual se correlacionó con medidas de longitud mediante regresiones lineales simples. Resultados: Las especies A. marta y A. caraca mostraron diferencias morfológicas estadísticamente significativas que permitieron confirmar estos taxones. El modelo potencial fue el que se ajustó mejor para mostrar las relaciones de la longitud total (LT) y el ancho de la cabeza (AC) con el peso seco (p<0.01). Conclusiones: Las relaciones entre las dimensiones corporales y el peso seco fueron altamente significativas, para A. marta el mejor ajuste fue con el AC y para A. caraca con la LT, con explicaciones del 86% y el 95% de la variación en la biomasa, respectivamente. Estas ecuaciones servirán como base para estudios de determinación de la biomasa en ninfas de Plecoptera.
Book
I Development in the Egg.- References.- II The Integument.- Properties of the cuticle.- Formation and shedding of the cuticle.- References.- III Growth.- Moulting.- Metamorphosis.- Determination of characters during post-embryonic development.- Regeneration.- Diapause.- References.- IV Muscular System and Locomotion.- Anatomy and histology.- Physiological properties of insect muscles.- Locomotion.- References.- V Nervous and Endocrine Systems.- Nervous system.- Visceral nervous system.- Endocrine system.- References.- VI Sense Organs: Vision.- Compound eye.- Simple eyes.- References.- VII Sense Organs: Mechanical and Chemical Senses.- Mechanical senses.- Hearing.- Chemical senses.- Temperature and humidity.- References.- VIII Behaviour.- Kinesis and related phenomena.- Orientation.- Co-ordinated behaviour.- References.- IX Respiration.- Tracheal system.- Development of the tracheal system.- Transport of oxygen to the tracheal endings.- Elimination of carbon dioxide.- Respiration of aquatic insects.- Respiration of endoparasitic insects.- Respiratory function of the blood.- Regulation of respiratory movements.- References.- X The Circulatory System and Associated Tissues.- Circulatory system.- Haemolymph.- Haemocytes.- Pericardial cells and so-called 'nephrocytes'.- Fat body.- Oenocytes.- Light-producing organs.- References.- XI Digestion and Nutrition.- Fore-gut.- Peritrophic membrane.- Mid-gut.- Hind-gut.- Secretions of the alimentary canal.- Digestion of some skeletal and other substances of plants and animals.- The role of lower organisms in digestion.- Nutrition.- References.- XII Excretion.- Urine.- Intermediary nitrogen metabolism.- Malpighian tubes.- Histophysiology of the Malpighian tubes.- Accessory functions of Malpighian tubes.- Malpighian tubes during moulting and metamorphosis.- Cephalic excretory organs and intestinal excretion.- Storage excretion.- References.- XIII Metabolism.- Chemical transformations.- Some chemical products of insects.- Pigment metabolism.- Respiratory metabolism.- References.- XIV Water and Temperature.- Water relations.- Temperature relations.- References.- XV Reproductive System.- Female reproductive system.- Male reproductive system.- Mating, impregnation and fertilization.- Some factors controlling fertility and fecundity.- Special modes of reproduction.- Sex determination.- Transmission of symbiotic micro-organisms.- References.- Index of Authors.- General Index.
Article
The regression equations for predicting dry mass from linear body dimensions presented in this study allow to calculate biomass of benthic macroinvertebrates from mountain streams of Central Europe. For the description of length-dry mass relationships, the power function y = a · xb or in a few cases the quadratic function y = a + b · x + c · x2 proved to be the best models. The relationships between linear body dimensions - e.g. body length (BL) vs. head capsule width (HW) - followed a simple linear model of the type y = a · x + b or again a quadratic one. Data of literature imply that the relationships derived from this study should be restricted to carbonate streams.