Survey of Twig-Inhabiting Coleoptera in Louisiana, USA

Abstract

Twigs are an understudied habitat utilized by many beetle species. Samples of twigs were collected in all six Level III Ecoregions in Louisiana during April and May 2013. The twigs were maintained in emergence chambers from which 942 specimens of adult Coleoptera were obtained, representing 31 families, 111 genera, and 128 species. Beetle richness and composition were not significantly different among ecoregions. Three species collected represent new state records: Pseudochoragus nitens (LeConte) (Anthribidae); Dryotribus mimeticus Horn (Curculionidae); and Narthecius grandiceps LeConte (Laemophloeidae). Four non-native species, all Curculionidae, were collected: Cyrtepistomus castaneus (Roelofs) (Entiminae); Cnestus mutilatus (Blandford) (Scolytinae); Xylosandrus crassiusculus (Motschulsky) (Scolytinae); and Xylosandrus germanus (Blandford) (Scolytinae).

Full text

The Coleopterists Bulletin, 70(3): 551–558. 2016.
SURVEY OF TWIG-INHABITING COLEOPTERA IN LOUISIANA, USA
MICHAEL L. FERRO
Clemson University Arthropod Collection
Department of Agricultural and Environmental Sciences
277 Poole Agricultural Center, Clemson University
Clemson, SC 29634-0310, U.S.A.
spongymesophyll@gmail.com
AND
NHU H. NGUYEN
Department of Environmental Science, Policy, and Management
University of California, Berkeley
130 Hilgard Way
Berkeley, CA 94720, U.S.A.
xerantheum@gmail.com
ABSTRACT
Twigs are an understudied habitat utilized by many beetle species. Samples of twigs were collected in all six Level III
Ecoregions in Louisiana during April and May 2013. The twigs were maintained in emergence chambers from
which 942 specimens of adult Coleoptera were obtained, representing 31 families, 111 genera, and 128 species. Beetle
richness and composition were not significantly different among ecoregions. Three species collected represent new
state records:Pseudochoragus nitens (LeConte) (Anthribidae); Dryotribus mimeticus Horn (Curculionidae); and
Narthecius grandiceps LeConte (Laemophloeidae). Four non-native species, all Curculionidae, were collected:
Cyrtepistomus castaneus (Roelofs) (Entiminae); Cnestus mutilatus (Blandford) (Scolytinae); Xylosandrus crassiusculus
(Motschulsky) (Scolytinae); and Xylosandrus germanus (Blandford) (Scolytinae).
Key Words: emergence chamber, beetles, fine woody debris, saproxylic, species richness
Of what use are twigs
but to sweep up litter
of fallen petals?
-Yosa Buson (translated by Harry Behn)
Twigs are more than a potential eye hazard to a
hiker, convenient kindling, or the bane of a subur-
banite’s lawn. Recent studies have revealed that
twigs and fine woody debris (FWD) are over-
looked habitats that support a rich beetle fauna.
A review of community-level surveys of North
American Coleoptera collected from FWD (up to
64 mm diameter) listed 83 species (Ferro et al.
2009). Collection of beetles emergent from mixed
hardwood FWD (25–70 mm diameter) in Great
Smoky Mountains National Park resulted in 1,544
specimens representing 162 species (Ferro et al.
2012). Two studies in Louisiana of twigs (14 ±
5 mm diameter) from Quercus falcata Michaux
(southern red oak, Fagaceae) resulted in the col-
lection of 414 beetles representing 35 species
(Ferro et al. 2009) and 39 beetles representing 21 spe-
cies (Ferro and Gimmel 2014).
Woody debris of all sizes is an important habi-
tat for many insects, especially Coleoptera (see
review in Ferro et al. 2012). Woody debris is gen-
erally divided into two size classes based on
diameter: coarse and fine. There are no standard-
ized size definitions. Commonly, the lower limit
of coarse woody debris (CWD) falls between a
diameter of 25–75 mm (Harmon et al. 1986), any-
thing smaller is considered FWD. Ferro et al.
(2012) defined FWD as woody debris with 25–
70 mm diameter and originating from trees. Ferro
et al. (2009) and Ferro and Gimmel (2014) con-
sistently used the term “twigs”to refer to woody
debris originating from trees that fell below 25 mm
diameter. In an effort to provide consistency, the
terms CWD, FWD, and twig will be used in this
study as defined above.
The purpose of this exploratory study was to
survey the beetle diversity within twigs through-
out various habitats in the state of Louisiana and
further develop the list of beetles that utilize the
twig habitat. A secondary purpose was to investi-
gate the community structure of twig-inhabiting
beetles among Louisiana ecoregions.
551

MATERIAL AND METHODS
North America is divided into a hierarchical
arrangement of ecoregions based on shared biotic
and abiotic characteristics (CEC 1997). Louisiana
contains two Level I, three Level II, and six
Level III Ecoregions (CEC 2006; Daigle et al. 2006)
(Table 1; Fig. 1). In an effort to thoroughly survey
twig beetle fauna within Louisiana, samples were
taken at each of the six Level III Ecoregions of
the state (Table 2). Specific sample sites were
chosen based on presence of forest cover and
availability of fine woody debris.
Samples consisted of dead twigs from hard-
wood trees, <25 mm diameter and broken into
<40 cm lengths. An effort was made to collect
twigs from a wide variety of tree species and cir-
cumstances, e.g., touching the ground, hanging
above the ground, attached to tree, dry, moist, lit-
tle decay, heavy decay. Twigs were collected dur-
ing April and May 2013 and immediately placed
into emergence chambers (see Ferro and Carlton
2011 for design). An April–May collection time
in Louisiana ensured that the twigs contained liv-
ing beetles presumably ready for a spring emer-
gence. An average of 180 twigs were placed in
each emergence chamber. A total of 36 samples
were taken at 19 locations, where each Level III
Ecoregion was represented by at least three loca-
tions (Table 2).
Emergence chambers were taken to Feliciana
Preserve (West Feliciana Parish, Louisiana) and
placed outdoors in a shaded, tree-covered area.
Beetles were allowed to emerge until 5 September
2013 (4–5 months) when all specimens were col-
lected. Propylene glycol was used as a killing and
preserving agent. Adult Coleoptera were pinned
or point-mounted as needed and labeled. Identifi-
cation to the lowest taxon level possible (typically
species) was performed with the appropriate taxo-
nomic literature and/or comparison with authorita-
tively identified reference specimens. All other
macroinvertebrates were retained, labeled, and pre-
served in 90% ethanol. Specimens are deposited in
the Louisiana State Arthropod Museum, Louisiana
State University, Baton Rouge, Louisiana.
Statistical comparisons of the community of
twig-inhabiting beetles were performed in R (R
Core Team 2015) with adonis in the vegan pack-
age (Okanen et al. 2005). Ecoregion, mean annual
temperature, annual precipitation, thermal line, and
hydric soil rating (US Geological Survey 1998)
were used as predictor variables for community dif-
ferences. Total richness was estimated using Chao-1
(Chao 1984).
RESULTS
In total, 942 specimens of adult Coleoptera
were obtained, representing 31 families, 111 gen-
era, and 128 species (or lowest taxonomic unit
when species could not be determined) (Table 3;
Appendix 1). Despite the size of this study with
36 samples at 19 locations (6,400+ twigs), few
species were recovered in abundance. Fifteen spe-
cies (12%) were represented by 10 or more speci-
mens, 56 (44%) were singletons, 70 (55%) were
collected from only a single ecoregion, 10 (8%)
were collected from five or more ecoregions, and
64 (50%) were collected from only one sample.
No significant differences in taxon richness
among ecoregions and no significant differences
among communities based on any of the predictor
variables were found (p> 0.05). Community sam-
pling was not saturated for any ecoregion (data
not shown). Chao-1 estimated a total richness of
228 species.
Table 1. Level I–III Ecoregions in Louisiana (CEC
2006). Numbers in parentheses correspond to Level III
Ecoregion designations from Daigle et al. (2006).
8.0 Eastern Temperate Forests
8.3 Southeastern USA Plains
8.3.5 (65) Southeastern Plains
8.3.6 (74) Mississippi Valley Loess Plains
8.3.7 (35) South Central Plains
8.5 Mississippi Alluvial And Southeast USA Coastal
Plains
8.5.2 (73) Mississippi Alluvial Plain
8.5.3 (75) Southern Coastal Plain
9.0 Great Plains
9.5 Texas-Louisiana Coastal Plain
9.5.1 (34) Western Gulf Coastal Plain
Fig. 1. Level III Ecoregions in Louisiana and col-
lection locations. See Table 2 for location information.
552 THE COLEOPTERISTS BULLETIN 70(3), 2016

DISCUSSION
Eleven species collected during this study over-
lapped with previous studies (Ferro et al. 2009;
Ferro and Gimmel 2014), bringing the total num-
ber of twig-inhabiting beetle species in Louisiana
to 167. Chao-1 estimated a total richness of 228 spe-
cies, showing that the sampling in this study only
found 64% of the expected number of beetles.
Finding that more species are expected was not
surprising as it is often the case in studies with
(what turned out to be) relatively small sam-
pling. However, the rarified dispersion of species
among the samples in this study was surprising. It
is in contrast to the high overlap of species among
twig bundles reported by Ferro et al. (2009),
which suggested a diverse but predictable com-
munity of beetles was associated with twigs.
Despite this contrast, the lack of community dif-
ferences among the various predictor variables
within Level III Ecoregions suggests that twig
communities are homogeneous and that twigs in
the broad sense are a uniform resource that are
home to a specific guild of beetles. However,
high species richness implies that variations within
the twig habitat (e.g., tree species, age, position,
Table 2. Localities and dates of twig collections. # = number on map (Fig. 1); numbers in parentheses correspond
to Level III Ecoregion designations from Daigle et al. (2006) (Table 1); (x2) = two samples taken from single location.
# Parish Location Latitude Longitude Date
(34) 1 Cameron Pevito Woods Sanctuary (x2) N 29.7576° W 93.6029° 20 April 2013
2 Jefferson Davis Jennings (x2) N 30.2376° W 92.6338° 20 April 2013
3 Vermilion Palmetto SP (x2) N 29.8662° W 92.1337 20 April 2013
(35) 4 Bossier Bayless Estate N 32.4496° W 93.8510° 11 May 2013
5 Bienville Big Cypress SP (x2) N 32.2552° W 92.9758° 12 May 2013
6 Catahoula Sicily Island WMA N 31.8118° W 91.7642° 14 April 2013
Sicily Island WMA N 31.8064° W 91.7729° 14 April 2013
7 Rapides Kincaid Recreation Area (x2) N 31.2672° W 92.6234° 11 May 2013
(65) 8 Tangipahoa Sandy Hollow WMA N 30.8016° W 90.3773° 4 April 2013
Sandy Hollow WMA N 30.8013° W 90.3800° 4 April 2013
9 Washington Bogue Chitto SP N 30.7803° W 90.1480° 4 April 2013
Bogue Chitto SP N 30.7796° W 90.1429° 4 April 2013
10 Washington 7 k W Bogalusa (x2) N 30.8569° W 89.9639° 4 April 2013
(73) 11 Concordia Bayou Cocodrie NWR (x2) N 31.5638° W 91.5601° 5 May 2013
12 Avoyelles Pomme de Terre WMA (x2) N 31.0280° W 91.8406° 14 April 2013
13 Iberville Sherburne WMA N 30.4193° W 91.6508° 14 April 2013
Sherburne WMA N 30.4248° W 91.6732° 14 April 2013
(74) 14 West Feliciana Tunica Hills WMA N 30.9377° W 91.5087° 4 May 2013
Tunica Hills WMA N 30.9322° W 91.5196° 4 May 2013
15 West Feliciana Feliciana Preserve N 30.7946° W 91.2537° 4 May 2013
Feliciana Preserve N 30.7956° W 91.2566° 4 May 2013
16 Livingston Tickfaw SP (x2) N 30.3827° W 90.6483° 4 May 2013
(75) 17 Tangipahoa East of Hammond (x2) N 30.5175° W 90.3733° 4 April 2013
18 Covington Fairview Riverside SP N 30.4111° W 90.1440° 4 April 2013
19 St. Tammany Honey Island WMA N 30.3741° W 89.6740° 4 April 2013
Honey Island WMA N 30.3879° W 89.6800° 4 April 2013
Table 3. Taxon richness by ecoregion. Numbers in parentheses correspond to Level III Ecoregion designations
from Daigle et al. (2006) (Table 1).
Ecoregion # families # genera # species
Western Gulf Coastal Plains (34) 19 37 39
South Central Plains (35) 17 40 42
Southeastern Plains (65) 16 34 34
Mississippi Alluvial Plains (73) 15 38 46
Mississippi Valley Loess Plains (74) 16 37 41
Southern Central Plain (75) 17 36 40
Total 31 111 128
553THE COLEOPTERISTS BULLETIN 70(3), 2016

season of death) result in a wide variety of niches,
and the life of a dead twig is more dynamic
than anticipated.
It is important to note that the lack of signifi-
cant differences in species richness or community
composition among Level III Ecoregions may be
influenced by the relatively small study size and
inability to saturate sampling. Additionally, this
and previous studies (Ferro et al. 2009; Ferro and
Gimmel 2014) are biased for species that over-
winter in twigs. However, even if these limitations
were overcome, the necessary fast life cycle and
vagility of most twig-inhabiting beetles may result
in a more or less homogenous community at
scales greater than Level III Ecoregions as seen
by Ferro et al. (2009). Additionally, aggregation
of twigs to a single location for emergence may
have influenced what species emerged, but that
seems unlikely due to the overall taxonomic diver-
sity observed.
Three species collected represent new state
records.Pseudochoragus nitens (LeConte)
(Anthribidae) (Fig. 2) was previously known only
from Massachusetts and Latimer Co., Oklahoma,
where it was collected by beating dead hickory
branches (Valentine 1991). Dryotribus mimeticus
Horn (Curculionidae) (Fig. 3) has a wide distribution,
but is only reported from Florida and South Carolina
in the USA and associated with driftwood along
beaches (Anderson 2002). Narthecius grandiceps
LeConte (Laemophloeidae) (Fig. 4) has only been
reported from Pennsylvania, Florida, and Latimer
Co., Oklahoma. It is a suspected predator of
Scolytinae (Curculionidae) and has been collected
from sweetgum and red oak (Thomas 1993).
Additionally, four non-native species, all Curcu-
lionidae, were collected: Cyrtepistomus castaneus
(Roelofs) (Entiminae); Cnestus mutilatus (Blandford)
(Scolytinae); Xylosandrus crassiusculus (Motschulsky)
(Scolytinae); and Xylosandrus germanus (Blandford)
Figs. 2–4. Species of new Louisiana state records. 2) Pseudochoragus nitens;3) Dryotribus mimeticus Horn;
4) Narthecius grandiceps.
554 THE COLEOPTERISTS BULLETIN 70(3), 2016

(Scolytinae). The scolytines are invasive species in
the USA. Cnestus mutilatus was reported damag-
ing plastic gas cans in Louisiana, presumably mis-
taking the container for desirable habitat (Carlton
and Bayless 2011).
Twigs are a convenient study system—they are
inexpensive to obtain, abundant, and easily col-
lected, transported, and stored. Furthermore, they
provide substrates in which beetles may be
“reared”in the field, and their transportability
allows for easy field manipulation to investigate
the ecology of twig-inhabiting beetles. The high
species richness and multiple guilds that twigs
harbor indicate that they may be useful for 1) util-
itarian studies (ecological indicators, impact, resto-
ration) as well as being 2) a worthy study system
unto themselves (fun). More general exploratory
studies and surveys are needed to better flesh out
the total diversity of beetles in twigs. However, the
findings of this research make it apparent that eco-
logical studies attempting to discover statistically
meaningful differences will have to be either much
larger or more specific in focus.
ACKNOWLEDGMENTS
We thank Andrew Flick, Jong-Seok Park,
Daniel Pessoa de Moura, and Brian Reily for help
in the field. Many thanks to Anthony Cognato
(Scolytinae), Matthew Gimmel (Phalacridae), Brittany
Owens (Staphylinidae), Jong-Seok Park (Staphylinidae),
Brian Reily (Cleridae), and Alexey Tishechkin
(Histeridae, Buprestidae) for help with species iden-
tifications. We thank Dorothy Prowell for allowing
the study to be conducted on her property.
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555THE COLEOPTERISTS BULLETIN 70(3), 2016

# Species
Level III Ecoregion
34 35 65 73 74 75
Anthribidae
1Choragus harrisii LeConte 1
2Euparius paganus Gyllenhal 1
3Eusphyrus walshi LeConte 1 1 4
4Euxenus punctatus LeConte 2 1
5Goniocloeus bimaculatus (Olivier) 1 2
6Piesocorynus mixtus LeConte 2 1
7Pseudochoragus nitens (LeConte) 1
Biphyllidae
8Diplocoelus rudis (LeConte) 1 2 1
Bostrichidae
9Lichenophanes bicornis (Weber) 1 1
Buprestidae
10 Acmaeodera tubulus (Fabricius) 1
11 Actenodes acornis (Say) 1
12 Agrilaxia flavimana (Gory) 2
13 Agrilus obsoletoguttatus Gory 1 1 1 2
14 Chrysobothris scitula Gory 2
15 Chrysobothris sexsignata (Say) 1
16 Xenorhipis brendeli LeConte 2 1
Carabidae
17 Mioptachys flavicauda (Say) 1 1 2 2 1
Cerambycidae
18 Astylopsis sexguttata (Say) 5
19 Curius dentatus Newman 2
20 Ecyrus dasycerus (Say) 9 2 3 8 1
21 Eupogonius pauper LeConte 1 1
22 Leptostylopsis planidorsus (LeConte) 2 1
23 Leptostylus transversus (Gyllenhal) 1 2
24 Lepturges angulatus (LeConte) 3
25 Lycochoriolaus lateralis (Olivier) 1
26 Methia necydalea (Fabricius) 1
27 Sternidius mimeticus (Casey) 1 1 1 1 5
28 Sternidius misellus (LeConte) 3 7
29 Sternidius punctatus Haldeman 1
30 Strangalia famelica Newman 1
Cerylonidae
31 Philothermus glabriculus LeConte 1
Ciidae
32 Orthocis longulus Dury 1
33 Orthocis punctatus (Mellie) 1 2
34 Orthocis transversatus (Kraus) 1 1
Cleridae
35 Chariessa pilosa (Forster) 1
36 Cymatodera wolcotti Barr 1
37 Neorthopleura thoracica (Say) 1
Corylophidae
38 Arthrolips sp. 1 1
39 Clypastraea lunata (LeConte) 2
40 Holopsis sp. 1 1 2 1
Cupedidae
41 Tenomerga cinerea (Say) 1
Curculionidae
42 Acalles clavatus (Say) 2 1 2 3
Continued on next page
APPENDIX 1
List of taxa and number of specimens collected from each ecoregion. Ecoregion designations are taken
from Daigle et al. (2006) (Table 1).
556 THE COLEOPTERISTS BULLETIN 70(3), 2016

Appendix 1. Continued.
# Species
Level III Ecoregion
34 35 65 73 74 75
43 Acalles minutissimus (LeConte) 3 2 9 1 1
44 Cnestus mutilatus (Blandford) 22
45 Cophes fallax (LeConte) 2 2 1 2
46 Cophes oblongus (LeConte) 1 1 1
47 Cophes obtentus (Herbst) 2
48 Cyrtepistomus castaneus (Roelofs) 1
49 Dryophthorus americanus (Bedel) 2 6 1 5 7
50 Dryotribus mimeticus Horn 1
51 Hylocurus langstoni (Blackman) 1
52 Hypothenemus eruditus Westwood 42 2 7 126 85 8
53 Hypothenemus interstitialus (Hopkins) 1
54 Hypothenemus sersatus (Eichhoff) 3 4 1 3
55 Laemosaccus nephele (Herbst) 1
56 Micracis swainei Blackman 6
57 Micracisella nanula (LeConte) 2
58 Phloeotribus texanus Schaeffer 1 1
59 Pityophthorus annectens LeConte 8 5
60 Pityophthorus pulicarius (Zimmermann) 7 1
61 Pseudopityophthorus asperulus (LeConte) 2 26 41
62 Pseudopityophthorus minutissimus (Zimmermann) 1
63 Thysanoes cf pallens Wood 4 8 9 9 3
64 Xylosandrus crassiusculus (Motschulsky) 1
65 Xylosandrus germanus (Blandford) 1
Elateridae
66 Idiotarsus errans (Horn) 2
67 Melanotus castanipes (Paykull) 1
Endomychidae
68 Anamorphus waltoni Blatchley 7 2
69 Aphorista vittata (Fabricius) 2 1 2
70 Clemmus minor (Crotch) 12823610
71 Stenotarsus blatchleyi Walton 1 1
Eucnemidae
72 Adelothyreus dejeani Bonvouloir 1
73 Dromaeolus teres (Horn) 1 1 1
74 Isorhipis nubila (Bonvouloir) 1 5
75 Microrhagus audax Horn 1
76 Nematodes atropos Say 1
Histeridae
77 Aeletes simplex (LeConte) 1 3
78 Bacanius (∼misellus LeConte) 1
79 Bacanius punctiformis (LeConte) 2 2 3 5
80 Epierus pulicarius Erichson 2 1 2
81 Paromalus seminulum Erichson 1
82 Platysoma leconti Marseul 1
Laemophloeidae
83 Narthecius grandiceps LeConte 1
Melandryidae
84 Microscapha clavicornis LeConte 11
Melyridae
85 Chaetocoelus setosus LeConte 1
Mordellidae
86 Glipostenoda ambusta (LeConte) 2
87 Mordella sp. 2
88 Mordellidae sp. 2
Nitidulidae
89 Stelidota octomaculata (Say) 1
Continued on next page
557THE COLEOPTERISTS BULLETIN 70(3), 2016

Appendix 1. Continued.
# Species
Level III Ecoregion
34 35 65 73 74 75
Phalacridae
90 Ochrolitus rubens (LeConte) 1 1
Ptiliidae
91 Pteryx sp. 1 1
Ptilodactylidae
92 Ptilodactyla angustata Horn 1
Ptinidae
93 Caenocara sp. 1
94 Calymmaderus nitidus (LeConte) 2 1
95 Cryptorama confusum White 1 4
96 Euceratocerus grandis White 3 1
97 Petalium sp. 1 4 2 1
98 Priobium sericeum (Say) 1
99 Tricorynus sp. 1 1 1
Salpingidae
100 Inopeplus immundus Reitter 1
Silvanidae
101 Silvanus muticus Sharp 1
Staphylinidae
102 Anotylus sp. 1
103 Athetini sp. 3 1 4
104 Belonuchus rufipennis (Fabricius) 4 1
105 Coproporus sp. 1
106 Dalmosella tenuis Casey 2 1
107 Edaphus americanus Puthz 1
108 Euconnus sp. 1 1 1
109 Eumicrota sp. 1
110 Eusphalerum sp. 1
111 Hesperus apicialis (Say) 2 2
112 Myrmecocephalus cingulatus (LeConte) 2
113 Pycnoglypta fracta (Casey) 4 1
114 Scydmaenidae sp. 1
115 Thinocharis sp. 1
116 Thoracophorus costalis (Erichson) 8 12 12 10 8 11
117 Toxidium gammaroides LeConte 1
Tenebrionidae
118 Isomira pulla (Melsheimer) 1
119 Platydema cyanescens Laporte and Brullé 1
120 Platydema flavipes (Fabricius) 1
121 Platydema ruficorne (Stürm) 1
122 Platydema sp. 1
123 Uloma mentalis Horn 1 1
Throscidae
124 Aulonothroscus convergens (Horn) 7 8 12 1 13 3
125 Throscidae sp. 1
Zopheridae
126 Endeitoma granulata (Say) 1
127 Paha laticollis (LeConte) 1
128 Pycnomerus sulcicollis (Germar) 1
558 THE COLEOPTERISTS BULLETIN 70(3), 2016

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Survey of Twig-Inhabiting Coleoptera in Louisiana, USA
Author(s): Michael L. Ferro and Nhu H. Nguyen
Source: The Coleopterists Bulletin, 70(3):551-558.
Published By: The Coleopterists Society
DOI: http://dx.doi.org/10.1649/0010-065X-70.3.551
URL: http://www.bioone.org/doi/full/10.1649/0010-065X-70.3.551
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