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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 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.
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 different 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
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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 different 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 different taxa may differ 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 different 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.
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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 coefficient 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 coefficient 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 coefficient 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 coefficients 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 coefficient 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
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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 coefficients 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 differ 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 differences
between the results obtained using different 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
¼coefficient of determination (*p50.01, **p50.001). n¼54.
230 B. Becker et al.
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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
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Some variations in length–dry mass relationships for the populations of the
same species, but from different locations, can be caused by physical–chemical
differences 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 affected 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 different 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), ‘Effects 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 Cliffs, NJ: Prentice Hall.
234 B. Becker et al.
Downloaded By: [Callisto, Marcos] At: 14:37 7 August 2009