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Analysis of the richness of Staphylinidae (Coleoptera) on different scales of a

sub-Andean rural landscape in Colombia

Article  in  Biodiversity and Conservation · June 2010

DOI: 10.1007/s10531-010-9812-2

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Biodivers Conserv
DOI 10.1007/s10531-010-9812-2

ORIGINAL PAPER

Analysis of the richness of Staphylinidae (Coleoptera)

on different scales of a sub-Andean rural landscape
in Colombia

Laura M. Vásquez-Vélez • Christian Bermúdez •

Patricia Chacón • Fabio H. Lozano-Zambrano

Received: 24 June 2009 / Accepted: 13 February 2010

Ó Springer Science+Business Media B.V. 2010

Abstract In the mid-watershed of the Nima River in the Colombian Central Cordillera,
the richness of staphylinids was estimated in association with 35 landscape elements
classified as mature forest, secondary forest, riparian corridors, pastures and cypress
plantations. For each element, two linear transects were traced, each one with six sample
units, consisting of a pitfall trap and a 1 m2 sample of leaf litter, processed in mini-Winkler
sacks. There were 2623 specimens grouped into 139 recognizable taxonomic units; 13
subfamilies were recorded: Pselaphinae, Aleocharinae, Paederinae, Staphylininae,
Tachyporinae, Osoriinae, Oxytelinae, Piestinae, Scaphidiinae, Euaesthetinae, Megalopsi-
diinae, Steninae and Omaliinae. The subfamilies with the greatest richness were Psela-
phinae (40 species), Aleocharinae (39) and Paederinae (22). The highest number of species
was found in the secondary forests (105 species), followed by the riparian corridors (98);
whereas the mature forest was the element with the fewest species (30). The diversity of
species components (a, b and c) was analyzed using the additive partition model based on
four spatial scales: sampling units, transects, elements and types of vegetative cover. The
elements and types of ground cover scales were the main contributors to total richness
percentages. The subfamilies Aleocharinae, Paederinae, Pselaphinae and Staphylininae had

L. M. Vásquez-Vélez (&)
Departamento de Biologı́a, Facultad de Artes y Ciencias, Universidad de Puerto Rico, Mayaguez,
PR 00681-9000, USA
e-mail: lauravasvel@gmail.com

C. Bermúdez
P. Chacón
Departamento de Biologı́a, Facultad de Ciencias Naturales y Exactas, Universidad del Valle,
Cali, A.A. 25360, Cali, Colombia
e-mail: christianax@gmail.com
P. Chacón
e-mail: pachacon@univalle.edu.co

F. H. Lozano-Zambrano
Instituto de Investigación de Recursos Biológicos Alexander von Humboldt,
Programa Biologı́a de la Conservación, CALLE 28A # 15-09, Bogotá DC, Colombia
e-mail: fhlozano@humboldt.org.co

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component diversity patterns that were equal to the overall assemblage. A significant
correlation of the richness of each subfamily with that of the assemblage was found.

Keywords Additive richness partition
Andean landscape
Beta diversity

Leaf litter
Staphylinidae

Introduction

Identification of the critical scales in which the ecological processes influence the diversity
of species is crucial for determining the appropriate levels where either habitat manage-
ment or ecological restoration should be implemented (Summerville et al. 2003; Chandy
et al. 2006). To monitor the effect of the changes in the environment and be able to design
successful conservation strategies (Moreno 2001, p. 21), it is necessary to estimate the
biological diversity according to different spatial scales: within natural and modified
communities (alpha diversity), exchanges among distinct communities or turnover (beta
diversity), and its contribution to the regional level (gamma diversity) (Whittaker 1960;
Halffter 2005).
Studies at the landscape level are particularly important for detecting and evaluating the
effects of anthropogenic activities on biodiversity (Halffter et al. 2001). To carry out this
type of study, indicator species should be chosen that reflect the changes in diversity
adequately under a variety of environmental conditions (Moreno and Halffter 2000).
McGeoch (1998) divides the indicators species into three categories: (1) environmental, (2)
ecological, and (3) biodiversity indicators.
The environmental and ecological indicators are used to detect changes in the environment,
while biodiversity indicators reflect the diversity of the overall biota (Rainio and Niemelä
2003), thus it is assumed that conservation-related decisions based on indicator species are
also appropriate for other organisms that live in the same area (Alonso 2000, p. 80).
Staphylinidae (Coleoptera) have been proposed as bioindicators or surrogates (Bohac
1999; Anderson and Ashe 2000; Pohl et al. 2007) given that they are abundant and diverse
(47,000 species); occupy a great variety of habitats worldwide; are easy to collect, given
that the majority of the species are found among leaf litter and fallen trunks, forming one of
the commonest groups of insects and ecologically the most important of the soil-related
fauna; and the determination of the majority of the subfamilies is relatively well known
(Bohac 1999; Navarrete-Heredia et al. 2002; Newton et al. 2005). In Colombia around 796
species and 230 genera of staphylinids are known, but some 5000 species are estimated,
placing this family in a taxonomic information gap (Newton et al. 2005). In the last decade
ecological studies of Staphylinidae from the Colombian Andes have been published,
revealing that in protected areas of the Eastern Cordillera there is a high exchange of
species in the altitudinal gradient from 700 to 2500 m (Gutiérrez and Ulloa-Chacón 2006).
In fragments of dry forest in the Cauca River geographic valley (900–1000 m altitude), a
positive relation was observed between richness of staphylinids and leaf litter moisture
(Garcı́a et al. 2001), as well as between richness and area of the fragments (Garcı́a and
Chacón de Ulloa 2005). Preliminary evaluations at the landscape level suggest that rich-
ness and regional diversity depend, to a great extent, on the habitats with greater plant
heterogeneity such as forests and riparian corridors (Bermúdez et al. 2006). In the Central
Cordillera, Sanabria et al. (2008) studied different land uses from 1000 to 1822 m, finding
that the richness of species in pastures with trees was comparable to what was found in
forests and stands of guadua (Guadua angustifolia). Recently, Gutiérrez-Chacón et al.

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(2009) found that there is a habitat specialitation for this family and it is related with
certain microhabitats characteristics that are found in riparian systems, such as humid litter,
mud and wet sand.
This work evaluated the alpha, beta and gamma components of the diversity of
Staphylinidae (Coleoptera) in the landscape of the mid-watershed of the Nima River, an
area that forms part of the zone surrounding Las Hermosas Natural National Park, located
on the western slope of the Central Cordillera. The purpose was to establish which spatial
scale contributed most to landscape richness and to find correlations between the richness
of the Staphylinidae subfamilies and that of the entire family in order to review its potential
use as a substitute for richness in studies focused on conservation.

Materials and methods

Study area

The mid-watershed of the Nima River is found on the western slope of the Central Cor-
dillera and belongs to the municipality of Palmira, Valle del Cauca Province (N3°350 5800 –
3°300 4000 W76°110 000 –76°100 5300 ), with an annual rainfall of 1000–1800 mm and a mean
temperature of 17°C (Mendoza et al. 2008). It corresponds to the natural zone of life found
from the premontane humid forest to the montane humid forest (Espinal 1967).
In this watershed a landscape area of 2500 ha, that spans within 1700–2000 m, was
delimited, where five elements of the landscape were selected in accordance with the
methodology for characterizing biodiversity in rural landscapes of the Conservation group
‘‘Paisajes Rurales’’ of the Alexander von Humboldt Institute (Mendoza et al. 2008): A patch
of mature forest ([100 ha) (Mf) with three replications; a patch of secondary forest
(between 3 and 100 ha) (Sf) with 13 replications; riparian corridor (Rc), which refers to
arboreal vegetation bordering watercourses with eight replications; pastures (Pa) with seven
replications; and cypress plantations (Cupressus lusitanica) (Cp) with four replications.
This gave a total of 35 replications of five types of vegetative cover in the landscape.

Sampling and taxonomic determination

The sampling was made from 22 July to 12 August 2006 and from 19 to 25 March of 2007,
covering the dry season for the area. The protocol implemented by Bermúdez et al. (2006)
was applied. In each replication two linear transects of 50 m were marked and separated by
a distance of not less than 50 m. Both transects included six sampling units, set up every
10 m and consisting of two methods of capture: Mini-Winkler sacks and pitfall traps,
placed parallel to the transect and separated by 10 m.
All the specimens collected were identified to the subfamily level and, where was
possible, to that of genus, using the keys of Navarrete-Heredia et al. (2002) and a revision
of Universidad del Valle Museum of Entomology collection. Operational taxonomic units
(OTUs) were used as substitutes for the species (Oliver and Beattie 1993, 1996).

Data analyses

The average efficiency of sampling, both at the level of the landscape elements and the
Staphylinidae subfamilies, was calculated based on two nonparametric estimators: Chao 1
and Jackknife 1 (Colwell 2005; Gotelli and Graves 1996). Box diagrams were used to

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visualize the variation of the richness among the replications, and the Kruskal–Wallis test
was applied to compare the richness among the types of vegetative cover. Rarefaction
curves were constructed based on the number of samples due to the fact that each element
had a different sampling intensity.
To determine the patterns of richness within the landscape, the hierarchical model of
additive partition of the components of diversity (c = a ? b) (Lande 1996) was applied,
where alpha is the average of the diversity within the habitats, beta is the exchange of
species among those habitats or turnover, and gamma is regional richness (Crist et al. 2003;
Gering et al. 2003). Within this study, the components were nested and represented by four
levels in the spatial scale: (1) sampling units composed by a 1 m2 sample of leaf litter and a
pitfall trap; (2) transects formed by six sample units; (3) landscape elements, a physical
unit that represents a type of vegetative cover; and (4) types of vegetative cover, a set of
elements of the landscape with the same type of ground cover. For example, the a1
component represents the diversity that exists within each sampling unit and b1 is the
exchange that occurs at that same level. The a of the next level is the sum of the com-
ponents of the lowest previous level. Regional diversity is represented by the following
formula: c = (a1 ? b1) ? (b2) ? (b3) ? (b4) (Fig. 1). Using the Partition program v.
2.0 (Veech and Crist 2007), a process of randomization (1000 times) was carried out to
generate null values for each of the components of diversity for each landscape scale in
order to compare statistically whether the observed values of richness were random or not.
The probabilities obtained from the randomization process were interpreted as the P-values
for the test of statistical significance. The same procedure was applied for the species that
were determined to be rare (percentage of abundance less than 1%) or common. Kol-
mogorov–Smirnov’s (Zar 1999) test for goodness of fit of nonparametric adjustments was
used to compare the components obtained for the total assemblage and each subfamily.
In addition, correlations were constructed between the subfamilies that had the greatest
richness of species and the family Staphylinidae for the levels that had significant dif-
ferences from what would be found at random in analyses based on additive partition, in
order to make a balance between the spatial levels at which the contribution to the land-
scape richness is higher, and the spatial level(s) where a real and effective management
plan can be made focused towards the conservation of the diversity. In each correlation the
Gamma (Landscape)

Alpha 4 (Types of vegetative coverage ) + Beta 4 (Exchange between types of vegetative

coverage )

Alpha 3 (Elements) + Beta 3 (Exchange between elements)

Alpha 2 (Transects) + Beta 2 (Exchange between transects)

Alpha 1 (Sampling Units) + Beta 1 (Exchange between sampling units)

Fig. 1 Relation among hierarchical levels in the additive partition model used for staphylinids in five types
of vegetative cover of the landscape in the Nima River mid-watershed (modified from Gering et al. 2003)

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subfamily analyzed was excluded from the rest of the family to ensure the independence of
both groups within the analyses and thereby observe how the richness of said subfamily
reflected the richness of the rest of the groups.

Results

General composition of staphylinids

A total of 2623 individuals were collected, grouped into 13 subfamilies, 60 genera and 139
OTU’s (Table 1). The subfamilies that contributed most to richness were Pselaphinae
(28%), Aleocharinae (25%), Paederinae (16%) and Staphylininae (11%). Considering the
entire Staphylinidae Family, an average sampling efficiency of 71% was obtained, but the
figure did not show an asymptotic tendency (Fig. 2). Looking at the subfamilies separately,
Aleocharinae had 75% efficiency, followed by Paederinae (73%), Pselaphinae (71%) and
Staphylininae (47%). The percentage of singletons (i.e. species represented by one indi-
vidual) was 30%, with the greatest number of singletons captures being for Pselaphinae
with 13 species, which represent 33% of the total of that subfamily for the study area;
Aleocharinae with 10 species (28%), Staphylininae with eight species (50%), Paederinae
with seven species (32%) and Scaphidiinae (50%) and Omaliinae with one species (100%).

Richness of staphylinids in five types of vegetative cover

The largest number of species (105) was observed in the secondary forest, followed by the
riparian corridor; while for the other three types of cover, which had similar values, the
number of species fell by one-third (Table 2). Significant differences (H = 15.93, gl = 4,
P \ 0.01) were found in the average richness by type of cover, highlighting a greater
variation in the riparian corridor in contrast with the mature forest (Fig. 2). The rarefaction
curve showed that upon interpolating the samples with those of the mature forest, which
was the cover with the least sampling intensity (36 sample units), the riparian corridor had
the greatest richness (68 species), followed by the secondary forest (53), the mature forest
(29), the cypress plantations (26) and pastures (23) (Fig. 3).

Analysis of the richness components

The mean richness (a4) within the different types of covers was significantly lower than
that found by chance (Table 3). The richness component that described the exchanges that
exist among the types of vegetative covers was the one that contributed most to regional
richness (b4 = 58.13%); and the value obtained was greater than that expected by chance.
The exchanges that exist among the landscape elements was the second component that
contributed most to regional richness, and this was significantly different from what would
have been expected by chance (b3 = 27.7%) (Fig. 4). Differences were found in the
patterns of richness between the rare and common species (Fig. 5). The species that were
defined as rare were the ones that had a pattern of richness similar to that of the total
assemblage (Kolmogorov–Smirnov: Rare: d = 0.4, P = 0.694; Common: d = 0.6,
P = 0.208). Exchange among the types of cover was the component that contributed most
to regional richness, but it was not significantly different from the expected random value.
The exchange at the level of the landscape elements was the one to which the commonest

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Table 1 Subfamilies of Staphylinidae (Coleoptera) associated with five landscape elements in the mid-
watershed of the Nima River (Valle del Cauca Province)
Subfamilies Tribe/Genus No. RTUs Mf Sf Rc Pa Cp Total

Aleocharinae Undetermined 36 24 344 214 52 10 644

Euaesthetinae Edaphus 1 – 7 8 1 – 16
Megalopsidiinae Megalopinus 2 2 5 3 – – 10
Omaliinae Phloeonomus 1 – – 1 – – 1
Osoriinae Aneocamptus 1 – 10 53 – – 63
Espeson 1 – 1 15 – – 16
Holotrochus 1 12 62 7 – 1 82
Mimogonus 1 1 3 – – – 4
Osorius 3 2 46 50 9 2 109
Oxytelinae Anotylus 2 – 63 101 24 3 191
Apocellus 1 – – 1 11 – 12
Dibelonetes 3 – 11 7 – – 18
Echiaster 1 1 18 20 – 1 40
Homaeotarsus 2 – 3 4 3 1 11
Lathrobium 1 – 1 – – – 1
LATHROBINA 2 1 8 2 – 1 12
Undetermined 1 – – – 1 – 1
Paederus 1 – 1 – – – 1
Palaminus 2 – 3 – 1 – 4
Ronetus 1 – 7 – – – 7
Scopaeus 1 – – 1 – – 1
Sphaeronum 1 – 1 1 – – 2
Suniocharis 1 2 1 2 – 1 6
Medon Gr. 3 – 7 9 7 – 23
Thinocharis 2 – 141 59 5 8 213
Piestinae Hypotelus 1 – 1 1 – – 2
Piestus 1 – 2 – – – 2
Pselaphinae S.T. JUBINI 13 4 82 148 1 17 252
SUPER TRIBU BATRISITAE 4 1 13 22 2 2 40
BRACHYGLUTINA 2 – 1 24 – – 25
Apharus 1 1 35 26 – – 62
Hamotus 1 – – 6 2 – 8
Listriophorus 2 – 8 9 – 3 20
TRICHONYCHINI 7 1 25 13 1 11 51
DIMERINI 1 – 9 17 – – 26
Rhexinia 1 2 1 3 – 1 7
Rhexius 1 – – 5 – – 5
TROGASTRINI 1 – 2 – – – 2
Phasmisus 1 – 1 1 – – 2
BYTHINOPLECTINA 2 – 4 5 – 1 10
Endytocera 1 – 1 – – – 1
GONIACERITAE 1 – – 1 – – 1
Leptoplectus 1 – 1 – – – 1

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Table 1 continued
Subfamilies Tribe/Genus No. RTUs Mf Sf Rc Pa Cp Total

Scaphidiinae Baeocera 1 3 9 3 – – 15
Scaphisoma 1 – – 1 – – 1
Staphylininae Chroaptomus 1 3 – 1 – – 4
Erichsonius 1 – 1 – – – 1
Microlinus 1 – – 1 – – 1
Heterothops 1 12 23 35 1 1 72
Diochus 2 – 3 2 2 – 7
Lithocharodes 2 1 16 22 3 1 43
Neohypnus 1 – – 1 – – 1
Paederominus 1 – – 10 – 1 11
Lissohypnus 1 – 1 – – – 1
Philonthus 3 1 6 14 10 – 31
Quedius 1 – – 1 – – 1
Tympanophorus 1 – – 1 – – 1
Steninae Stenus 1 – 1 6 – – 7
Tachyporinae Bryoporus 3 1 12 6 – – 19
Coproporus 2 – 193 155 22 3 390
Sepedophilus 1 2 2 8 – – 12
Total 139 77 1,196 1,106 158 69 2,623

Tribes are typed in capitalized letter and genus in italic letters

Mf mature forest, Sf secondary forest, Rc riparian corridor, Pa pastures, Cp cypress plantation

species contributed most. For the rare species, the highest levels (b3 and b4) of the land-
scape scale contributed most of the 91.25% of regional richness (Table 3).
On comparing the patterns of richness for each subfamily with those of the total
assemblage, no significant differences were found in the contribution of each component of
richness (Kolmogorov–Smirnov: Pselaphinae d = 0.2, P = 0.999; Paederinae d = 0.2,
P = 0.999; Staphylininae d = 0.2, P = 0.999; Aleocharinae d = 0.2, P = 0.999) (Fig. 4).
When the richness of the subfamilies was compared with the total richness of the
family, positive and significant correlations were found at the level of the landscape
elements in Pselaphinae (r = 0.68, gl = 33, P \ 0.001), Paederinae (r = 0.71, gl = 33,
P \ 0.001), Staphylininae (r = 0.73, gl = 33, P \ 0.001) and Aleocharinae (r = 0.64,
gl = 33, P \ 0.001). At the level of the vegetative cover, significant regressions were
found in Pselaphinae, Paederinae and Aleocharinae, but not for Staphylininae (Fig. 5).

Discussion

Distribution of the richness of staphylinids in the landscape

Regional richness received the greatest contribution from the exchange of species found at
the level of types of ground cover and elements, revealing the existence of an effect of the
differences in vegetative cover and probably, in the physical characteristics of each ele-
ment (Fragment size, shape of the perimeter, isolation, type of edge) on the composition of

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A 160

140

120
Richness (S) 100

80
Obs
60
CI 95%
40

20

0
0 200 400 600 800

Number of samples

B 120

100
Richness (S)

80

60 Mf
Sf
40
Rc

20 Pa
Cp
0
0 50 100 150 200 250 300
Number of Samples

Fig. 2 a Accumulation curve for the entire landscape of the Nima River mid-watershed. b Accumulation
curve for species in the five types of vegetative cover. Mf mature forest, Sf secondary forest, Rc riparian
corridor, Pa pastures, Cp cypress plantation

Table 2 Richness, abundance and sampling efficiency for the assemblage of the staphylinids in five types
of vegetative cover in the mid-watershed of the Nima River
Values Mf Sf Rc Pa Cp

Abundance (No. of individuals) 77 1,199 1,125 158 69

Richness observed (No. of species) 30 105 98 34 32
Average estimated richness (Chao 1 and Jackknife 1) 45 150 141 48 60
Average sampling efficiency (%) 66.4 70.2 69.6 70.8 53.3

Mf mature forest, Sf secondary forest, Rc riparian corridor, Pa pastures, Cp cypress plantation

the staphylinids. Similar results have been found in other group of ground beetles (Cara-
bidae), where the sizes of patches changes the species composition, species number and/or
species abundance (Rainio and Niemelä 2003). According to Niemelä (2001) area effects,

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60

50

Mean Richness
40

30

20

10

0
Mf Sf Rc Pa Pc
Elements

Fig. 3 Comparison of the richness of staphylinids in five types of vegetative cover of the landscape in the
Nima River mid-watershed. Mf mature forest, Sf secondary forest, Rc riparian corridor, Pa pastures,
Cp cypress plantation

Table 3 Average richness of Staphylinidae (Coleoptera) found for the different levels of sampling and the
significance of each value; greater than that expected at random (?); there are no differences from what
would be expected at random (NS); less than expected at random (-)
Level Total Rare species Common species Pselaphinae Paederinae Aleocharinae Staphylininae

a1 3 (-) 1 (-) 1.8 (-) 1 (-) 1 (-) 1 (-) 1 (-)

b1 9 (-) 5 (-) 4 (-) 2 (-) 2 (?) 2 (-) 1 (NS)
b2 7 (-) 4 (-) 2 (NS) 2 (-) 1 (NS) 2 (NS) 1 (NS)
b3 39 (?) 33 (?) 6 (?) 11 (?) 6 (?) 10 (?) 5 (?)
b4 81 (?) 77 (NS) 4 (NS) 24 (NS) 13 (-) 25 (NS) 9 (-)

edge effects, shape of the fragment and habitat connectivity have impacted the ground
beetle assemblage. These characteristics appear to be very important for structuring the
richness of the assemblage of Staphylinidae in the zone. On dividing the assemblage
between rare and common species, it is evident that those characteristics affect only the
least abundant species. Of both groups, the pattern most similar to that of the total
assemblage was that of the rare species, in which the lowest scales were the ones that
contribute least to the richness; and the highest scales were the ones that contribute most
(Fig. 4). This could be due to the fact that these rare species, as they were defined,
represent 87% of the species found in this sampling; or because in reality they are truly
important within the assemblage from an ecological standpoint. It is not yet clear, as
defined by Gering et al. (2003), whether these rare species are the result of the sampling
design, whether they are biologically significant or whether they are really rare.
High percentages of singletons and rare species have been found in other studies of
Coleoptera with a greater sampling extension (Gering et al. 2003; Bermúdez et al. 2006).
However, this work did not have a very large extension and although the efficiency of
sampling was relatively high, the percentage of singletons continued to be high (30%).

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Fig. 4 Percentage of the total richness of Staphylinidae explained by the diversity components for four
spatial scales: sampling units, transects, elements and types of vegetative cover

Fig. 5 Correlation among the richest subfamilies of Staphylinidae and the total assemblage excluding the
subfamilies given, taking into the account the spatial scale of landscape elements. From the upper left to the
lower right: Paederinae richness versus the richness of the rest of the Staphylinidae family, Stahylininae
richness versus the richness of the rest of the Staphylinidae family, Pselaphinae richness versus the richness of
the rest of the Staphylinidae family and Aleocharinae richness versus the richness of the rest of the Staphylinidae
family

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This could be due to the megadiversity of the family and/or to the use of OTUs, which
could be masking the reality of the identities and the real number of species that exists in
this area. The common or more abundant species could have broader ranges of tolerance to
the differences in plant composition and characteristics of the elements, which would mean
that these characteristics do not have much influence on the structure of their richness,
abundance and composition. This can be seen in the contribution that each landscape scale
makes to their richness, where the exchanges that exist among the types of cover do not
differ from what would be expected by chance, suggesting that for this group in particular,
the difference in plant composition is not a determining factor.
The diversity of staphylinids found in this work corresponds to 68% of the subfamilies
and 17.5% of the species recorded for Colombia (Newton et al. 2005). The subfamilies
Pselaphinae, Aleocharinae, Paederinae and Staphylininae, which had the highest richness,
are also the ones that have the largest number of species worldwide (Navarrete-Heredia
et al. 2002) and have been the most recorded in previous research. The pattern observed in
this study with respect to the richness of species in these subfamilies is similar to that found
by Gutiérrez and Ulloa-Chacón (2006), where Pselaphinae and Aleocharinae represented
more than 50% of the species collected. However, the richness of Pselaphinae was not high
in proportion to the other subfamilies in productive systems such as those of the Cauca
River geographic valley (Bermúdez et al. 2006), where the crop growing of sugarcane
dominates, and in cattle-raising zones of Quindı́o, where there were no collections (San-
abria et al. 2008).
In general, the individuals of Pselaphinae, Aleocharinae, Paederinae and Staphylininae
are predators; within each subfamily some generalist species can be found that frequent
leaf litter, decomposing trunks, carrion, excrement, remains of fruits, caves, banks of
bodies of water, etc. There are also genera that are highly specialized as to the type of prey
that they eat (e.g. some Pselaphinae) or to the environments where they live, such as those
that are myrmecophilous (Aleocharinae, Paederinae and Staphylininae) (Navarrete-Here-
dia et al. 2002).
With hyperdiverse groups such as insects, it is difficult to obtain the complete inventory
of the species that inhabit a given area (Jiménez-Valverde and Hortal 2003), even more so
with Staphylinidae, which is the second most diverse family of the Order Coleoptera
(Navarrete-Heredia et al. 2002). The efficiency of average sampling depends, above all, on
the common species that are introduced more rapidly into the inventory (Jiménez-Valverde
and Hortal 2003). It is to be expected that groups such as Aleocharinae, where there are a
large number of common species and that exploit more habitats (Navarrete-Heredia et al.
2002; Majka and Klimaszewski 2008), would be collected more than those of subfamilies
with rarer species. It should be noted that two subfamilies (Omaliinae and Scaphidiinae)
were collected that had not been recorded in earlier studies (Garcı́a et al. 2001; Garcı́a and
Chacón de Ulloa 2005; Bermúdez et al. 2006; Gutiérrez and Ulloa-Chacón 2006; Sanabria
et al. 2008). From the subfamily Omaliinae, one individual from the genus Phloeonomus
(Table 1) was found, which although it had been recorded in Colombia (Newton et al.
2005), an exact location was not given. The species of this subfamily are characterized by
having highly varied feeding habits: predators, saprophagous or pollen feeders; they are
found mainly in leaf litter, under the bark of fallen trunks and in flowers (feeding on pollen).
The individuals from the genus Phloeonomus appear to be associated primarily with flowers
of palms (Arecaceae) (Navarrete-Heredia et al. 2002). In the subfamily Scaphidiinae, which
until recently was considered a family separate from Staphylinidae, two genera were found:
Baeocera and Scaphisoma (Table 1). The individuals of this subfamily feed on spores and
hyphae of different groups of fungi (Navarrete-Heredia et al. 2002).

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The most favorable vegetative covers in the landscape for harboring a high degree of
richness of staphylinids were the secondary forest and the riparian corridor, independent of
the number of samples (Fig. 2). The mature forest, which could be an element with
vegetative cover appropriate for the staphylinid fauna, did not contribute to a high number
of species, which could be due to the low number of replications for this element in the
landscape. After interpolating the number of samples of all the vegetative covers in the
rarefaction test, the mature forest would have the third highest richness. These results can
be explained by two approaches: First, the Intermediate Disturbance Hypothesis (Connel
1978; Roxburgh et al. 2004), where the mature forests is the habitat with low disturbance,
the pastures and cypress plantation are the habitat with high frequency of disturbance and
the secondary forest with the intermediate disturbance frequencies. In the mature forest the
better competitor excludes the good disperser and this can explain the lower richness of
this type of cover; in the pastures and cypress plantations, the better competitor but poorer
disperser cannot persist, so the richness also decrease. At intermediate disturbance fre-
quencies, like the one found in the secondary forest, there is a parameter zone where both
species can coexist indefinitely, with the competitively inferior species occupying the
recently disturbed sides, what contributes the increase of richness (Roxburgh et al. 2004).
The second approaches, is the fragments size. The difference between the size of mature
forests and the secondary forests, might explain the difference between their richness. It
has been observed that the high number of species of ground beetles found in small-sized
(0.5–3 ha) fragments can be explained by the invasion of species from the surroundings of
the fragments (Halme and Niemelä 1993).
The riparian forest richness can be due to the similarity of the flora composition to the
secondary forest, sharing almost the 50% of the species (IAVH 2008), although the
abundant of each species is different, showing differences in the dominance patterns. On
the other hand, this habitat provide environments and conditions that are especially suitable
for staphylinids such as decomposing trunks and more moisture in the leaf litter (Garcı́a
et al. 2001; Navarrete-Heredia et al. 2002). According with Gutiérrez-Chacón et al. (2009),
the riparian rove beetles are found almost exclusively close to the water0 s edge or in very
wet habitats (mud, wet sand) in all their developmental stages.
Cypress plantations have been found to be strongly favorable to organisms that exploit
habitats similar to those of staphylinids, such as ants (Jiménez and Lozano-Zambrano
2005); however in the study zone this type of cover is an important source of economic
exploitation, where the intensity of their management (fertilization, fumigation and cut-
ting) can have an unfavorable effect on the fauna in the soil. Despite the foregoing, the
cypress plantations conserve greater richness than the pastures. This may be due to the fact
that the canopy and the layer of leaf litter create more favorable conditions than those
found in pastures.

Relations of richness within Staphylinidae

The proportions of singletons found in the whole assemblage and in each one of the richest
subfamilies are similar. This can explain the similarity between the patterns of alpha and
beta richness of each subfamilies and the total assemblage. The rare species could be the
factor that determines the observed pattern of alpha and beta richness for the Staphylinidae
family in this landscape.
For these same subfamilies, however, the exchanges at the spatial scale of the landscape
elements was significantly higher that what would be expected at random; whereas at the
spatial scale of the vegetative cover, the exchange was either not significant (Aleocharinae

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and Pselaphinae) or was lower than what would be expected at random (Paederinae and
Staphylininae) (Table 3). At both spatial scales, the richness of these subfamilies and that
of the other Staphylinidae were closely related, where the change in the richness of one
group conditioned the richness of the other. This supports what was found with the additive
partition analyses, where the patterns of richness of the total assemblage are reflected in
each of these subfamilies. The foregoing reiterates the importance of the different char-
acteristics of all the elements that form the landscape, making them distinctive for the
staphylinids.
These results indicate that staphylinids can be considered as good surrogates for
evaluating species richness, almost within the family. It was shown by the correlation
between the richest subfamilies and the rest of family that certain groups can reflect the
richness of others and may be used in the construction of rapid biodiversity surveys. The
patterns found in the partition analysis also support this idea (Fig. 4). Here are proposed
some groups of staphylinids as indicators or surrogates of the whole assemblage of the
Staphylinidae family. Despite the complexity of the family taxonomy, we can focus on
some of the subfamilies or groups in which there is a better resolution in the identity of the
genera and maybe there is a better possibility to achieve a specific determination. The
subfamily that can accomplish those characteristics is Staphylininae because they are
the third subfamily with more species of the entire family (Navarrete-Heredia et al. 2002),
the biology and morphological characteristic are relatively well known, they are generally
big in size, are one of the most conspicuous groups inside the family and they can be
collected with more types of leaf litter sampling methods than other subfamilies (Psela-
phinae) (LV and CB pers. Obs.).
In the tropics there are few studies about beetles in general used as indicators (Pearson
and Cassola 1992; Rainio and Niemelä 2003) and particularly, the staphylinids have been
poorly used as indicator species in this area (Anderson and Ashe 2000). Due to the loss
rate of the Andean forest the best way of knowing the biodiversity in these places is
through the use of the rapid evaluation on the remaining habitats. The use of the
Staphylininae subfamily as a richness indicator is a good way to accomplish this.
Nonetheless the use of morphospecies is questioned; some works have obtained a low
percentage of error between the number of real species and the number of morphospecies
(Oliver and Beattie 1993, 1996; Gering et al. 2003). One of the characteristics of a good
indicator is that its taxonomic resolution most be clear (Alonso 2000, p. 80; Rainio and
Niemelä 2003), but due to the low percentage of error that the use of morphospecies can
represent. The use of staphylinids as surrogates can be supported by the use of
morphospecies.

Contributions for conservation

The result of the analyses of the richness components for this landscape, where the
exchanges of species between the types of cover and the elements are what contribute most
to regional richness, provide another argument for proposing conservation schemes from
the perspective of the rural landscapes for the Colombian Andes. More than 70% of the
Andes in the country correspond to rural landscape zones, and the conservation of the
patches of forest alone will not guarantee the conservation of the regional diversity of
species. It is important to propose conservation schemes that value and maintain the
diversity of habitat and the heterogeneity of the landscapes given that these attributes are
those that are promoting and increasing the diversity.

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Acknowledgments The authors are grateful to the Instituto de Investigation of Recursos Biológicos
Alexander von Humboldt for providing the financing for this project; to the communities in the villages of
Tenjo and La Quisquina in the municipality of Palmira, Valle del Cauca. Gratitude is also expressed to
Catalina Gutiérrez for the help provided in identifying the specimens, to Professor Deborah Letourneau for
her comments on the manuscript and to Margaret K. Thayer for her valuable comments on the biology of
Staphylinidae. The authors also wish to thank the Royal Embassy of the Netherlands, the Global Envi-
ronmental Facility and the World Bank for financing the project ‘‘Conservation and Sustainable Use of the
Biodiversity in the Colombian Andes,’’ within the framework of which this research was conducted. Authors
are also thankful for the valuable comments from the reviewers of this paper.

References

Alonso L (2000) Ants as indicators of diversity. In: Agosti D, Majer JD, Alonso LE, Schultz TR (eds) Ants:
standard methods for measuring and monitoring biodiversity. Smithsonian, Washington, DC, pp 80–98
Anderson RS, Ashe JS (2000) Leaf litter inhabiting beetles as surrogates for establishing priorities for
conservation of selected tropical montane cloud forest in Honduras, Central America (Coleoptera:
Staphylinidae and Curculionidae). Biodivers Conserv 9:617–653
Bermúdez C, Gutiérrez C, Chacón P (2006) Estimación de la diversidad de coleópteros Staphylinidae en el
paisaje del bosque seco tropical. In: Libro de Resúmens II Congreso Colombiano de Zoologı́a, Santa
Marta, Colombia, p 407
Bohac J (1999) Staphylinid beetles as bioindicators. Agric Ecosyst Environ 74:357–372
Chandy S, Gibson DJ, Robertson PA (2006) Additive partitioning of diversity across hierarchical spatial
scales in a forested landscape. J Appl Ecol 43:792–801
Colwell RK (2005) EstimateS (v. 7.5b1), Copyright RK Colwell: http://viceroy.eeb.uconn.edu/estimates
Connel JH (1978) Diversity in tropical rain forest and coral reefs. Science 199:1302–1310
Crist TO, Veech JA, Gering JC, Summerville KS (2003) Partitioning species diversity across landscapes and
regions: a hierarchical analysis of a, b and c diversity. Am Nat 12(6):734–743
Espinal S (1967) Apuntes sobre ecologı́a colombiana. Universidad del Valle, Departamento de Biologı́a,
Cali, Colombia, p 45
Garcı́a R, Chacón de Ulloa P (2005) Estafilı́nidos (Coleoptera: Staphylinidae) en fragmentos de bosque seco
del valle geográfico del rı́o Cauca. Rev Colomb Entomol 31(1):43–50
Garcı́a R, Armbrecht I, Ulloa-Chacón P (2001) Staphylinidae (Coleoptera): Composición y mirmecofilia en
bosques secos relictuales de Colombia. Folia Entomol Mex 40(1):1–10
Gering JC, Crist TO, Veech JA (2003) Additive partitioning of species diversity across multiple spatial
scales: implications for regional conservation of diversity. Conserv Biol 17(2):488–499
Gotelli N, Graves GR (1996) Null models in ecology. Smithsonian Institution Press, Washington, DC, p 368
Gutiérrez C, Ulloa-Chacón P (2006) Composición de estafilı́nidos (Coleoptera: Staphylinidae) asociados a
hojarasca en tres localidades de la Cordillera Oriental de Colombia. Folia Entomol Mex 45(2):69–81
Gutiérrez-Chacón C, Zúñiga MC, Van Bodegom PM, Chará J, Giraldo LP (2009) Rove beetles (Coleoptera:
Sataphylinidae) in Neotropical riverine landscapes: characterising their distribution. Insect Conserv
Divers 2:106–115
Halffter G (2005) Towards a culture of biodiversity conservation. Acta Zool Mex (n.s.) 21(2):133–153
Halffter G, Moreno CE, Pineda EO (2001) Manual para evaluación de la biodiversidad en reservas de la
biosfera. MandT-Manuales y Tesis SEA, vol 2. Zaragoza, p 80
Halme E, Niemelä J (1993) Carabid beetles in fragments of coniferous forest. Ann Zool Fenn 30:17–30
Instituto Alexander von Humboldt (2008) Diversidad, composición y estructura de la vegetación de un
paisaje rural ganadero de la cuenca media del rio Nima, Palmira. Grupo Conservación de la Biodiv-
ersidad en Paisajes Rurales, Instituto Alexander von Humboldt, 22 p
Jiménez E, Lozano-Zambrano FH (2005) Diversidad alfa, beta y gama de hormigas de suelo en paisajes
rurales ganaderos de los Andes centrales colombianos. In: Memorias V Coloquio de Insectos Sociales
IUSSI-Sección Bolivariana, Cali, Colombia, pp 41–56
Jiménez-Valverde A, Hortal J (2003) Las curvas de acumulación de especies y la necesidad de evaluar la
calidad de los inventarios biológicos. Rev Iber Aracnol 8:151–161
Lande R (1996) Statistics and partitioning of species diversity, and similarity among multiple communities.
Oikos 76:5–13
Majka CG, Klimaszewski J (2008) New records of Canadian Aleocharinae (Coleoptera: Staphylinidae). In:
Majka CG, Klimaszewski J (eds) Biodiversity, biosystematics and ecology of Canadian Coleoptera.
ZooKeys 2, pp 85–114

123
Biodivers Conserv

McGeoch M (1998) The selection, testing and application of terrestrial insects as bioindicators. Biol Rev
73:181–201
Mendoza JE, Jiménez E, Lozano-Zambrano FH, Caycedo PC, Renjifo LM (2008) Identificación de ele-
mentos del paisaje prioritarios para la conservación de biodiversidad en paisajes rurales de los Andes
Centrales de Colombia. In: Harvey C, Saénz J (eds) Evaluación y conservación de biodiversidad en
paisajes fragmentados de Meso América. Santo Domingo de Heredia: Instituto Nacional de Biodiv-
ersidad, INBio, pp 251–285
Moreno CE (2001) Métodos para medir la biodiversidad. M and T-Manuales y Tesis SEA, vol 1. Zaragosa,
p 84
Moreno C, Halffter G (2000) Assessing the completeness of bat biodiversity inventories using species
accumulation curves. J Appl Ecol 37:149–158
Navarrete-Heredia JL, Newton AF, Thayer MK, Ashe JS, Chandler DS (2002) Guı́a ilustrada para los
géneros de Staphylinidae (Coleoptera) de México. Universidad de Guadalajara and CONABIO,
Guadalajara, Mexico, p 401
Newton AF, Gutiérrez-Chacón C, Chandler DS (2005) Checklist of the Staphylinidae (Coleoptera) of
Colombia. Biota Colomb 6(1):1–72
Niemelä J (2001) Carabid beetles (Coleoptera, Carabidae) and hábitat fragmentation: a review. Eur J
Entomol 98:127–132
Oliver I, Beattie AJ (1993) A possible method for the rapid assessment of biodiversity. Conserv Biol 7:562–
568
Oliver I, Beattie AJ (1996) Invertebrate morphospecies as surrogates for species: a case study. Conserv Biol
10:99–109
Pearson DL, Cassola F (1992) World-wide species richness patterns of tiger beetles (Coleoptera: Cicin-
delidae): indicator taxon for biodiversity and conservation studies. Conserv Biol 6:376–391
Pohl GR, Langor DW, Spence JR (2007) Rove beetles and ground beetles (Coleoptera: Staphylinidae,
Carabidae) as indicators of harvest and regeneration practices in western Canadian foothills forests.
Biol Conserv 37(1):294–307
Rainio J, Niemelä J (2003) Ground beetles (Coleoptera: Carabidae) as bioindicators. Biodivers Conserv
12:487–506
Roxburgh SH, Shea K, Wilson JB (2004) The intermediate disturbance hypothesis: patch dynamics and
mechanisms of species coexistence. Ecology 85(2):359–371
Sanabria MC, Armbrecht I, Gutiérrez C (2008) Diversidad de estafilı́nidos (Coleoptera: Staphylinidae) en
cinco sistemas productivos de los Andes. Rev Colomb Entomol 34(2):217–223
Summerville KS, Boulware MJ, Veech JA (2003) Spatial variation in species diversity and composition of
forest Lepidoptera in eastern deciduous forests of North America. Conserv Biol 17(4):1045–1057
Veech JA, Crist TO (2007) Partition v. 2.0. Software for the additive partitioning of species diversity
Whittaker RH (1960) Vegetation of the Siskiyou Mountains, Oregon and California. Ecol Monogr
30(3):279–338
Zar JH (1999) Biostatistical analysis. Prentice-Hall, New Jersey, p 417

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