Longhorned Beetles (Coleoptera: Cerambycidae and Disteniidae) Collected in the Canopy of a Dry Tropical Lowland Forest in Panama

Abstract

This study characterizes the abundance and richness of longhorned beetles (Coleoptera: Cerambycidae and Disteniidae)
collected in the canopy of a dry tropical lowland forest in Panama. The nine tree species sampled included Enterolobium
cyclocarpum (Jacq.) Griseb. (Fabaceae), Pseudobombax septenatum (Jacq.) Dugand (Malvaceae), Luehea seemannii Triana
& Planch. (Malvaceae), Cordia alliodora (Ruiz. & Pav.) Oken (Boraginaceae), Castilla elastica Sessé ex Cerv. (Moraceae),
Anacardium excelsum (Bertero & Balb. ex Kunth) Skeels (Anacardiaceae), Cecropia peltata L. (Urticaceae), Zuelania
guidonia (Sw.) Britton & Millsp. (Flacourtiaceae) and Ficus insipida Willd. (Moraceae). A total of 227 specimens of
Cerambycidae and Disteniidae were collected. The most abundant species was Trestonia pulcherrima Dillon and Dillon,
1946 (23%), followed by Compsibidion vanum (Thomson, 1867) (16%). The greatest abundance and richness of beetles
were recorded in the canopy of L. seemannii (90 individuals = 39%). The presence of these beetles in the forest canopy is
associated with their general preference for large trees, highly complex foliar canopy architecture, and the availability of
resources for the development and feeding of larvae and adults. Synchronization with plant phenology may explain the
greater abundance and richness observed in L. seemannii.

Full text

651
The Coleopterists Bulletin, 75(3): 651–660. 2021.



Universidad de Panamá, Centro Regional Universitario de Colón
Escuela de Biología, Departamento de Zoología, PANAMA
and

Universidad de Panamá, PANAMA
alfredo.lanusa@up.ac.pa




and


alpizar@sfe.go.cr


Programa Centroamericano de Maestría en Entomología
Universidad de Panamá, PANAMA
and
Universidad de Panamá, Departamento de Ciencias Ambientales

enrique.medianero@up.ac.pa

A
This study characterizes the abundance and richness of longhorned beetles (Coleoptera: Cerambycidae and Disteniidae)
collected in the canopy of a dry tropical lowland forest in Panama. The nine tree species sampled included Enterolobium
cyclocarpum (Jacq.)   Pseudobombax septenatum (Jacq.) Dugand (Malvaceae), Luehea seemannii Triana
& Planch. (Malvaceae), Cordia alliodora  (Boraginaceae), Castilla elastica 
Anacardium excelsum     Cecropia peltata   Zuelania
guidonia   and Ficus insipida Willd. (Moraceae). A total of 227 specimens of
Cerambycidae and Disteniidae were collected. The most abundant species was Trestonia pulcherrima Dillon and Dillon,
Compsibidion vanum   
were recorded in the canopy of L. seemannii
   
        
greater abundance and richness observed in L. seemannii.


I
The forest canopy is defined as the crown of each
tree in the forest, including leaves, branches and
epiphytes, and is the functional interface between

et al. 2003). The role of the can
opy is paramount in sustaining high levels of
diversity, as the canopy is an important repository
of biological richness (Ballerio and Wagner 2005;

652 
Castaño et al. 2006; Charles and Basset 2005;
Novotny and Basset 2005; Novotny et al
Numerous insect species spend most of their lives
in the treetops in both tropical and temperate for
ests, and they provide a variety of food resources
for the organisms that occupy the canopy. The

them important energy assimilators, leading to
higher rates of primary production (e.g., flowers,
fruits, and seeds) and consequently larger amounts
of resources for herbivorous insects, including

et al. 2020;
Novotny et al. 2003).
Many species of beetles in the canopy are habitat
specialists (Barrios 2012), whereas others are gen
eralists that use the canopy only temporarily
(Castaño et al. 2006; Winchester et al
Cerambycidae and Disteniidae, the canopy pro
vides an array of resources for their broad feeding
habits (Meng et al. 2013; Vance et al. 2003); con
sequently, they are among the most common beetle
groups collected during tropical canopy surveys
     
These beetles are mainly associated with living and
    
et alet al. 2003);
such resources not only support Cerambycidae in
et al. 2017) but also
     
 et al     
many species of longhorned beetles is poor, espe
cially species inhabiting inaccessible environ
ments, such as the upper layers of trees, as they are
et al.

beetle fauna of the canopy as well as the relation
ship between them and the canopy in tropical for
 
et alet al. 2003).
et al. (2006) in the tropical
       
    
have provided some insight into beetle species in
      
et al
  
beetles for different forest strata as well as the mi
croclimatic conditions determining the composi
tion of the longhorn beetle community. The

cerambycid and disteniid species associated with
the canopy of nine tree species representative of
tropical dry forest; 2) determine whether there is
an association between specific longhorned beetle
species and trees.
MM
  The study was conducted in

     
     
study site is classified as a lowland tropical semide
ciduous forest and is located at the transition be
tween tropical humid forest and the premontane dry



ha; its rainy season runs from May to November,
and its dry season from December to April. The
upper canopy is 25–30 m, with emergent trees

is within the limits of Panama City; along with
the Parque Nacional Camino de Cruces and

    
     
Barrios 2017).
A construction crane installed in this forest by the
    

       

2 of forest (Wright and Colley


including trees and lianas (Ødegaard 2000). The
dominant species in the canopy are Luehea seeman-
nii Triana & Planch. (Malvaceae), Anacardium
excelsum      
(Anacardiaceae), Ficus insipida Willd. (Moraceae),
Castilla elastica 
Cordia alliodora    
(Boraginaceae). At the site where the crane is lo
       
(Medianero et al.
et al. (2017), the area
of the forest covered by the crane has the character
istics of deciduous forest.
A Malaise trap modified for canopy studies was
       
were made with organza fabric and were 66.5 cm
long and 60 cm high; a polyethylene collection
       
highest end.
A trap was placed in each of the following nine
tree species: Enterolobium cyclocarpum (Jacq.)
  Pseudobombax septenatum
(Jacq.) Dugand (Bombacaceae), L. seemannii, C.
alliodora, C. elastica, A. excelsum, Cecropia peltata
Zuelania guidonia 
Millsp.   F. insipida. These

  653
species were selected because they are the tallest
local trees. An additional trap was placed in trees of
L. seemannii, A. excelsum and F. insipida because
two or more specimens of each of these species were
located within the area of the crane.
 
          

because the high evaporation rates in the canopy
might damage samples with longer sampling in
tervals. During each sampling event, the contents
of the collection bottles were collected in a con
tainer, and samples were labeled by noting the tree
species, trap number, and date of collection. The
collection bottles were refilled with alcohol and

tion period. The samples were then transported to
one of the laboratories of the Central American

of Panama. Provisional species identifications

graphic catalog for Cerambycidae and Disteniidae.
     
parison with available specimens in the collections




     
      
  
     
Maes et al    et al. (2013),
     
et al. (2020).
    
each of the sampled tree species were placed in
   
generated a list of individuals by species collected.
The normality of the data was assessed by the


in the richness and abundance of longhorn beetles
present in the different tree species studied.
    
to determine if there was a relationship between
the species of longhorned beetles and the identity
of the tree species where traps were placed. The



A total of 227 adult Cerambycidae and Disteniidae
were collected, encompassing 37 species in 31 gen
era, 17 tribes and two subfamilies of Cerambycidae,


    
only five species were represented by 10 or more
individuals (n
of these, Trestonia pulcherrima Dillon and Dillon,

while Compsibidion vanum  
Chlorida festiva  Obrium cordi-
colle    Aerenea impetiginosa

all collected specimens (Table 1).
     H
p
differences in the number of Cerambycidae col
lected among the sampled plants. However, the
largest number of species was collected in traps
placed in trees of L. seemanniiA.
excelsum (52 specimens), C. elastica (35 speci
mens), F. insipida  E. cyclo-
carpum
were collected in the trap placed in C. peltata. The
     
placed in L. seemannii and A. excelsum were T.
pulcherrima and C. vanum; in the traps located in
F. insipida and C. elastica  
species was C. vanum
The simple correspondence analysis indicated
that there was a relationship between cerambycid
species and plant species where traps were locat
ed (X2p
variance (inertia) was 1.63; however, the analysis

was the month when the largest number of cer
ambycid specimens was collected in the canopy

 Composition of species, number and per
centage of Cerambycidae and Disteniidae specimens
associated with the canopy of nine tree species in the
MNP.
n



Total  100
  63
Two specimens 6 20
≥ 10 specimens 5 16.7

Total 227 100
Most abundant species
(Trestonia pulcherrima)
52 
 132 71.7

 
  Tree species
Enterolobium cyclocarpum (Entcyc), Pseudobombax septenatum (Pseusep), Luehea seemannii Cordia alliodora (Corall), Castilla elastica (Casela), Anacardium excelsum
Zuelania guidonia (Zuegui) and Ficus insipida




       



Chlorida festiva   1  3 1 3  

Coleoxestia sanguinipes  2   2   

Megacyllene angulata  1  1    

Anelaphus misellus  31  1  1

Compsa macra  3 1 1 1     
Compsibidion vanum  37 11 6 10 2  
Neocompsa squalida  2     2 

Obrium cordicolle    3 6 1 3 1  

Euderces cleriformis  11     

Trachyderes succinctus succinctus  1     1 


Atrypanius exilis  1   1   
Atrypanius haldemani    2 1 1    
Lagocheirus araneiformis ypsilon  51  1 2 1 
Leptostylus decipiens     5    
Leptostylus leucopygus  1     1 
Leptostylus x-griseus  1     1 
Lepturges (Chaeturges) inscriptus  32 1     
Lepturges (L.) laeteguttatus  11     
Lepturges (L.) sexvittatus  11     
Nealcidion privatum  1    1  
Stenolis inclusa  62 1 1 1 1
Urgleptes tumidicollis    6  3  
(continued)

  655

Oreodera albilatera  21   1 

Adetus bacillarius  11     
Dorcasta dasycera (Erichson in 1    1  

Colobothea dispersa  21 1     

Aerenea brunnea  5 1 3 1     
Aerenea impetiginosa  10 3 2 3 2   

Desmiphora (D.) niveocincta  1  1    
Estola ignobilis  32  1  
Estoloides prolongata  1   1   

Taeniotes scalatus  1    1  

Oncideres argentata  1  1    
Trestonia pulcherrima  52 27 12 5  
Venustus zeteki      1 2 1  

Ataxia fulvifrons    2  2  1
Epectasis juncea  1  1    


Cupecuara turnbowi  31 1   1 
Elytrimitatrix (Grossifemora) geniculata  11     
 227 3  52  35  2 2
   26  11 15 12 2 2
 (Continued.)




       

656 
Number of Cerambycidae and Disteniidae individuals collected with Malaise traps in nine tree species in
       
25 species. No specimens were collected in the trap placed in Cecropia peltata; this tree species is therefore not rep
resented in the graph.


     
Disteniidae have a preference for trees of the family
Malvaceae, followed by Moraceae and Anacardiaceae.
  
  

et al. 2006; Medianero et
al  
(2017) indicated that the Malaise trap method is ef
fective for collecting unique species of Cerambycidae

  657
compared with other traditional methods used for the
same purpose. Most of the sampled beetles belonged
        
group in the tropics, larger than Cerambycinae

belonged to the tribe Acanthocinini, which contains
adults that are mostly active at high temperatures and
et al
According to Pringle et al. (2012), larger trees
attract a greater abundance and richness of herbiv
orous insects than smaller trees because they have

more insects per unit leaf area, such as the high
diversity and abundance of cerambycids in this
study from Luehea, Anacardium, Ficus, Castilla and
Enterolobium.  
mental role in the feeding habits of both larvae and
      
canopy branches, and adults consume nutrient
sources, such as pollen or leaves, or simply do not
consume food at all (Carvalho et al 
  
ously or seasonally available resources in the can
      

of cerambycids (Novotny et al. 2003).
The abundance and richness of longhorned bee
tles associated with the L. seemannii canopy can be

chronization with plant phenology. The flowering
of L. seemannii occurs between November and
June; however, from December to January, the pro
duction of flower buds increases (Condit et al.
2011), which would support a greater number of
associated individuals and species (Table 2) as well
as a greater overall seasonal abundance during this
      et al.
(2017) support this idea, as the richness and diver
sity of Cerambycidae were synchronized with the

The low richness and abundance observed in the
other tree species in this study were puzzling. The
low herbivory rates in C. alliodora might stem from
the presence of biological defenses associated with

eggs and larvae of longhorned beetles in the canopy,
similar to observations made by Way et al
in Portugal, and Palmer and Brody (2013) in Acacia

has low foliar herbivory, which might reflect the


posed to generalist species (Mitter et al

groups, the structure of trees, the quantity, distribu
tion, and availability of resources, and phenology can
influence the composition of longhorned beetles in
habiting the canopy. Although challenging, canopy
studies have become increasingly important for ob
      
understood forest layer. Nontraditional methods for
collecting Cerambycidae and Disteniidae might have

However, given the effectiveness of these methods,
they provide a potential alternative approach for elu
  
nity assembly.
A
    

      

       
     


 Ecology and diversi
ty of canopy associated Ceratocanthidae (Insecta:
    
rainforest [pp. 125–132]. In: African Biodiver
     




secta: Coleoptera) del dosel de árboles tropicales.

 A composite interception trap for sam
pling arthropods in tree canopies. Australian
     doi.

   Invertebrates in the canopy of trop
ical rain forests How much do we really
     

      
      
and moisture continuum with conservatism and
   In  
Diversity, Ecology and Conservation (M. D. Uly
    

     Host utilization

    Palame (Coleop
   
 
 

  A photographic catalog of the Ce
rambycidae of the world. New World Ceramby
cidae catalog. bezbycids.com/byciddb/wdefault.

     
New species of Cerambycidae from Panama, with

 
new distribution records (Coleoptera: Cerambyci

 Review of the
genus Cupecuara (Coleoptera: Disteniidae). Acta
Entomologica Musei Nationalis Pragae 60(2):

   

de una parcela de bosque en el Parque Natural


  Medium
  
      



    
        


       
    
Chrysomelidae) in two forest types in Panama.


        
of the Cerambycidae and Disteniidae of North


   


  Trees of
Panama and Costa Rica. Princeton University

    Comparison of
the species richness and abundance of Ceram
    
tera: Curculionidae) captured in aerial Malaise
traps with and without a bottom collector. The
    

    Revision of the Neotropical Acan
thocinini (Coleoptera: Cerambycidae). The genus
Lagocheirus. Bulletin of the British Museum of
Natural History (Entomology) 6: 137–166.

patterns, trophic structure and richness of Colop
tera in the tropical arboreal ecosystem: The fauna
of the tree Luehea seemanii Triana and Planch
in the Canal Zone of Panama. The Coleopterists

      
Estructura de la comunidad del orden Diptera
       

   Review of the
genus Coleoxestia 
tral America, part I (Coleoptera, Cerambycidae).

   Review of the
genus Coleoxestia 
tral America, Part II (Coleoptera, Cerambycidae).

    Introduction: Cano
py arthropod research in Europe [pp. 13–20]. In:

     
Nuremberg, 576 pp.

tral American species of Colobothea 
(Coleoptera: Cerambycidae). The Coleopterists

  A review of
the genus Euderces   
ambycidae: Tillomorphini). Proceedings of the


(Coleoptera, Cerambycidae) collected in Mo
       
      

      Novas es
  Cometes   
    
   

  Biol

(Coleoptera: Cerambycidae). Journal of Entomol

      
     Dead
      
  
     

     Es
   

sidad y abundancia. Boletín del Museo de Ento

     
    
beetles go up when rains come down. Biotropica

        
      
    Catálogo ilustrado de los
     
cidae. Revista Nicaraguense de Entomologia 70:

        
 
 Catálogo ilustrado de los

Disteniinae. Revista Nicaraguense de Entomolo

 Tribo Ceram
Coleoxestia [pp.
 In  
 

       
 Cerambycidae (Coleoptera) collected
 
zon forest canopy and the description of four new
species. Acta Amazonica 36(2): 265–272. doi.


  
       
     
  Composición de la co
munidad de artrópodos que habitan en el dosel


        
 Tree diversity mediates the distribution of
longhorn beetles (Coleoptera: Cerambycidae)
   
       

       
The phylogenetic study of adaptive zones:
    
     

       
      

without erect setae on elytra; synonymies, trans
ferences, revalidation, and notes on genera with


 


    

 
  
      
predatory guilds of forest canopy arthropods along
     





   
Achievements and horizons in canopy science.
  

    
     



in the tropical dry forest. Neotropical Entomolo


        
insect herbivores in tropical forests. Proceed
       
   

     Herbi
vore communities and their food resources [pp.
In  


     


     
  
on woody plants from one hectare of a lowland


   The relative importance of trees
versus lianas as hosts for phytophagous beetles
(Coleoptera) in tropical forests. Journal of Bioge



tles in the tropical tree Brosimum utile (Moraceae):
    
   


     
       
  
Biodiversity meets the atmosphere: A global view
  

        Enough is
   
on herbivory, growth, and reproduction in an
     

         
Plant defense, herbivory, and the growth of Cor-
dia alliodora trees and their symbiotic Azteca
     

       Descrip
tions, transferences and synonymies in Ameri
can Monochamini (Coleoptera, Cerambycidae,
 Ham-
matoderus and Taeniotes.  


   
Distenia
 
  

      
ware system), version 7. www. statistica.soft
ware.informer.com/7.0 (accessed 20 July 2021).
 Zonas de Vida. Una Base Ecológica Para
     
        
Naciones Unidas para la Agricultura y la Ali

 
Community composition of longhorned beetles
(Coleoptera: Cerambycidae) in the canopy and
understorey of sugar maple and white pine stands

   

      
   
     
Phoracantha semipunctata (Coleoptera: Cer
ambycidae) in Portugal. Bulletin of Entomolog
    


660 

     
   
    

      Accessing the Can
opy. Assessment of Biological Diversity and Mi

United Nations Environmental Program, Nairobi,


leoptera: Phytophagoidea). Academia de Ciencias

(Received 30 December 2020; accepted 30 July 2021.
Publication date 7 December 2021.)