A Key to the Forensically important sap beetles

(Coleoptera: Nitidulidae) of North America

Gareth S. Powell1,2 and Lauren M. Weidner3*
1
Department of Biology, Brigham Young University, Provo, UT, USA
2
Florida State Collection of Arthropods, Division of Plant Industry, Florida Department of Agriculture
and Consumer Services, Gainesville, FL, USA
3
School of Mathematical and Natural Sciences, New College of Interdisciplinary Arts and Sciences,
Arizona State University– West Campus, Glendale, AZ, USA

*corresponding author: [Link]@[Link]

Editor:
Section labels:

Abstract

The forensically relevant taxa of the family Nitidulidae (Insecta: Coleoptera) are examined and diagnosed.
Specifically, the genera Nitidula and Omosita are known to be forensically important based on the shared
feeding behavior on carrion. Eight species are reported from North America (excluding Mexico), including
a new country record for Omosita funesta Reitter. Distribution maps for each species were generated from
collection records compiled from 25 institutions with major holdings from the USA and Canada. The
validity of Omosita nearctica Kirejtshuk was upheld by genetic distances of COI DNA barcode data from
four species of Omosita. Finally, a dichotomous key is provided for the Nitidula and Omosita species,
supplemented with dorsal habitus photographs of each species.

Keywords
Omosita, Nitidula, carrion, Nitidulinae
Nitidulidae, or sap beetles, are small (1 mm– Mortem Interval (mPMI) of remains in a more
15mm long) oval to elongate beetles with clubbed advanced stage of decay.
antennae with a global distribution (Jelínek et al.
2010). There are estimated to be approximately
Omosita colon was the most frequent species of
4000 species worldwide, with approximately 200
this family collected (when identified down to
of these species occurring within the United
species) from decomposition studies (Reed 1958,
States and Canada (Habeck 2002). Members of
Rodriguez 1982, Watson and Carlton 2003,
this family are much more ecologically diverse
2005). Shubeck et al. (1981) noted that O. colon
than the common name might suggest. This
was the dominant beetle species collected (35% of
family contains species that are fungivorous,
all beetles collected, 99.71% of all nitidulids)
frugivorous, anthophilous, predatory, and
from baited traps in New Jersey. However,
necrophagous in addition to some exhibiting
nitidulids are not always recorded during
saprophagy (Parsons 1943, Powell et al. 2020).
decomposition studies, or they are identified only
Necrophagy is exhibited by two genera, Nitidula
to family (McLeod 2015) or genus (Abdell et al.
and Omosita (Parsons 1943). Despite this well–
1982, Monthei 2009). The further into
known feeding strategy, they are often overlooked
decomposition remains are, the more challenging
in forensic systems, possibly due to their small
it becomes to provide an accurate mPMI
size (not collected) or not easily identifiable
estimation. Understanding the species
(identified to family).
composition of the insect communities in these
advanced stages of decomposition is the first step
Nitidulids have been collected during the later to determine which insects should be studied in
stages of decomposition across all seasons greater detail. Here we present an overview of the
(Anderson and VanLaerhoven 1996, Watson and forensically relevant species of Nitidulidae for
Carlton 2005) from a variety of vertebrate North America north of Mexico with distribution
remains including dogs (Reed 1958), swine maps for each species and a diagnostic key in an
(Adair and Kondratieff 1996), rats (DeJong and attempt to increase the utilization of these
Hoback 2006) various wildlife (bears, alligators, important species in forensic studies.
deer, turtles, and gulls; Abdell et al. 1982, Lord
and Burger 1984, Watson and Carlton 2003, Materials and Methods
2005) and humans (Rodriguez 1982). Even Specimens examined
though they are frequently encountered, little
work has been completed on life histories in a The following public and private collections
forensic context. Zanetti et al. (2013) provided an contain specimens that were physically examined
overview of larval morphology and development for the present study.
of Nitidula carnaria (Schaller) and Wang et al.
ARCC Andrew R. Cline Private Collection
(2020) examined the development of Omosita
(Sacramento, CA)
colon (L.) at a range of temperatures and provides
insight to their thermal maximum threshold, AEC Arthur Evans Private Collection
which aids in estimating the minimum Post– (Richmond, VA)
BYU Monte L. Bean Museum, Brigham Young TAMC Texas A&M University Insect Collection
University (Provo, UT) (College Station, TX)

CNC Canadian National Insect Collection UAF University of Alaska Insect Collection
(Ottawa, CAN) (Fairbanks, AK)

CMN Canadian Museum of Nature (Ottawa, UCFC University of Central Florida Collection
CAN) (Orlando, FL)

CUAC Clemson University Arthropod Collection UMSP University of Minnesota Insect Collection
(Clemson, SC) (St. Paul, MN)

CSCA California States Collection of Arthropods USNM United States National Museum,
(Sacramento, CA) Smithsonian (Washington, D.C.)

FMNH Field Museum of Natural History WIRC University of Wisconsin Research

(Chicago, IL) Collection (Madison, WI)

FSCA Florida State Collection of Arthropods Distributions

(Gainesville, FL)
State or province level presence data were
GSPC Gareth S. Powell Private Collection recorded for the USA and Canada from more than
(Lafayette, IN) 4,000 physical specimens from the collections
listed above. Additional records were also
INHS Illinois Natural History Survey compiled from published literature (Connell
(Champaign, IL) 1984, Dodge 1937, Downie and Arnett 1996,
KESC Kyle E. Schnepp Private Collection Dury 1902, Majka and Cline 2006, Powell 2015,
(Gainesville, FL) Price and Young 2006, Vogt 1950, Williams et al.
1997). Coarse (state/province–level) distribution
MPM Milwaukee Public Museum (Milwaukee, maps were produced in QGIS Essen v2.14.0
WI) (QGIS Development Team, 2020).

MSU A.J. Cook Collection, Michigan State Key development

University (East Lansing, MI)
Couplets were in part adapted and combined from
NHM Natural History Museum (London, UK) several previous works (Parsons 1943; Audisio
1993) and then modified and illustrated here.
NZAC New Zealand Arthropod Collection
Additional diagnostic tools have recently been
(Auckland, NZ)
presented for the Palearctic (Lee and Lee, 2015)
PERC Purdue Entomological Research and for South Africa (Williams et al., 2021).
Collection (West Lafayette, IN) Character images were acquired using a Leica
DFC450 camera mounted onto a M165C
PSUC Frost Museum, Penn State University stereomicroscope or a Vision Digital Passport
(State College, PA) imaging system with a Canon EOS 6D camera
with a 65mm lens.
RMBC R. Michael Brattain Private Collection
(Lafayette, IN) DNA–based species confirmation
Representative DNA barcodes (cytochrome c of this practical application to forensic
oxidase subunit 1 “CO1”) were acquired from entomology in North America.
NCBI GenBank for four species of Omosita
(Genbank Accession numbers given after each Barcode confirmation of Omosita neartica
taxon); O. colon 1 (MT653618), O. colon 2 DNA barcodes were used to generate a genetic
(KM452482), O. colon 3 (KM441201), O. colon distance matrix (Table 1) to assess the validity of
4 (KM446224), O. depressa (KU916617), O. four species of Omosita. Genetic similarity is
depressa 2 (KM439454), O. discoidea 1 given as a percentage (Table 1). Two species, O.
(KU912774), O. discoidea 2 (HQ953668), O. discoidea and O. nearctica, were each represented
discoidea 3 (KM445991), O. discoidea 4 by multiple individuals and those individuals
(KU919455) and O. nearctica 1 (MT371766), O. were shown to be >98% identical across the
nearctica 2 (MG054067), O. nearctica 3 barcode region of CO1.
(MG058703). Nitidula bipunctata (KU918404)
was included as an outgroup. Sequences were Genetic similarity between the four species of
aligned following default conditions using Omosita was shown to be 84.91-91.21% 85.52–
MAFFT v.7.45 (Katoh and Standley 2013) 89.97%. The historically confused O. colon and
implemented in Geneious Prime v.2021.0.1 O. nearctica exhibit the upper end of that range,
(Kearse et al. 2012). Genetic distances were also with 89.97% similarity. While barcodes alone are
generated in Geneious Prime. a contentious method of defining or diagnosing
species (Brower 2006, Rubinoff et al. 2006), they
can be valuable tools and, in this case, clearly
Results and Discussion support the validity of O. nearctica as
morphologically described by Kirejtshuk (1987)
Taxonomic notes

Nitidula nigra Schaeffer, 1911, is treated here as

a synonym of N. rufipes. Audisio (1993) argued
for this nomenclatural change based on a
combination of shared characters (i.e., elytra with
narrowly flattened margins, lacking clear elytral
patterning, and aedeagal structure) and lack of
reliable morphological differences between the
two taxa. Omosita colon (Linnaeus, 1758) was by
far the most prevalent name used for the
morphospecies that is the most commonly
collected in the United States and Canada with a
lighter apex of the elytra. Kirejtshuk (1987)
described O. nearctica, to which most of these
records are referring but stated that a few
confirmed specimens of O. colon were seen from
parts of Canada. None of those records were able
to be confirmed for the present work and remain
rare historic records that are not included as part
Table 1. Genetic similarity (%) matrix between several species of Omosita based on the barcode region of COI.

N_bipunctata O_colon_1 O_colon_2 O_colon_3 O_colon_4 O_colon_2 O_depressa_2 O_depressa O_discoidea_2 O_discoidea_3 O_discoidea_
N_bipunctata X
O_colon_1 87.08 X
O_colon_2 85.71 87.99 X
O_colon_3 85.87 87.99 99.85 X
O_colon_4 85.71 87.99 100.00 99.85 X
O_colon_2 85.87 87.99 99.85 100.00 99.85 X
O_depressa_2 84.35 86.32 86.17 86.02 86.17 86.02 X
O_depressa 84.80 86.63 86.32 86.47 86.32 86.47 99.24 X
O_discoidea_2 86.17 88.75 88.30 88.30 88.30 88.30 86.47 86.47 X
O_discoidea_3 86.17 88.75 88.30 88.30 88.30 88.30 86.47 86.47 100.00 X
O_discoidea_4 86.17 88.75 88.30 88.30 88.30 88.30 86.47 86.47 100.00 100.00 X
O_discoidea_5 86.17 88.75 88.30 88.30 88.30 88.30 86.47 86.47 100.00 100.00 100.00
O_discoidea_1 86.17 88.75 88.30 88.30 88.30 88.30 86.47 86.47 100.00 100.00 100.00
O_nearctica_3 85.02 91.21 90.23 90.23 90.23 90.23 87.30 87.62 90.55 90.55 90.55
O_nearctica_2 85.11 89.97 89.06 89.21 89.06 89.21 84.95 85.56 87.84 87.84 87.84
O_nearctica_1 85.21 89.94 89.02 89.18 89.02 89.18 84.91 85.52 87.96 87.96 87.96
Range Expansions (1943) reported N. flavomaculata from California
and a second introduction near Washington, D.C.
General distributions are given for each species We now report N. flavomaculata from over a
(Fig 2A–E, Fig. 3A-C). Many of the included taxa dozen states/provinces. Omosita funesta is added
are not thought to be native to North America and to the fauna of the United States as two specimens
have instead been moved by humans (Parsons, were studied from southern Texas that clearly
1943). This means that several of the known match this name (deposited in FMNH). The
distributions are disjunct and should be species was known from northern Mexico prior to
considered tentative. Nitidula flavomaculata for this study and so this only represents a minor
example was not known to occur in the area prior range expansion in published distribution for this
to the middle of the 20th century but we now report species.
from a much larger range. Parsons

Fig. 2. North American north of Mexico distributions for each species of forensically important Nitidulidae. (A)
Nitidula bipunctata, (B) Nitidula carinaria, (C) Nitidula flavomaculata, (D) Nitidula rufipes, (E) Nitidula ziczac.
Fig. 3. North American north of Mexico distributions for each species of forensically important Nitidulidae. (A)
Omosita discoidea, (B) Omosita funesta, (C) Omosita neartica.

Checklist of Nitidula and Omosita for North 1a. Labrum bilobed (Fig.4A) and the presence of
America north of Mexico paramedial pronotal depressions………..2
(Omosita)
Nitidula bipunctata (Linnaeus, 1758)
b. Labrum emarginate (Fig. 4B) and a lack of
Nitidula carnaria (Schaller, 1783) paramedial pronotal depressions.…...4 (Nitidula)
Nitidula flavomaculata (Rossi, 1790) 2a. Elytra nearly parallel, pronotum with long,
Nitidula rufipes (Linnaeus, 1767) golden setae along basal third of margin………

Nitidula ziczac Say, 1824 …………………..……………..Omosita funesta

Omosita discoidea (Fabricius, 1775) b. Elytra narrowing posteriorly, with nearly

indistinct silver setae…………………….…….3
Omosita funesta Reitter, 1873 new country
record 3a. Base of elytra distinctly darker than apex
………………………………...Omosita neartica
Omosita neartica Kirejtshuk, 1987
b. Apex of elytra distinctly darker than base, disc
Dorsal habitus photos (Figs. 5-6) are provided for of pronotum also darker …...…Omosita discoidea
all eight species to assist the key below.
4a. Elytra unicolorous ………….Nitidula rufipes

b. Elytra with distinct maculations ……………5

Key to adult Nitidula and Omosita in the USA
and Canada 5a. Each elytron with single pale spot (often
orange) ……………………...Nitidula bipunctata
 b. Elytra lacking well defined single spots……6 Anthony Davies (CNC), Andrew Deans (PSUC),
Arthur Evans, Michael Ferro (CUAC), Michael
6a. Pronotum broadly explanate with margins Geiser (NHM), François Génier (CMN), Richard
lighter……………………Nitidula flavomaculata Leschen (NZAC), Eugenio Nearns (PERC),
b. Pronotum not explanate…. ………………7 Crystal Maier (FMNH), Thomas McElrath
(INHS), Sandor Kelly (UCFC), Edward Riley
7a. Body length shorter, 1.5–3.0mm…………… (TAMU), Kyle Schnepp (FSCA), Floyd Shockley
(USNM), Derek Sikes (UAF), Paul Skelley
……………….……….........… Nitidula carnaria
(FSCA), Robin Tompson (UMSP), Dan Young
b. Body length longer, 3.0– (WIRC), Jennifer Zaspel (MPM). We thank
5.0mm..…………………….…..Nitidula ziczac Natalie Saxton for assistance taking dorsal habitus
photos and Laura Sutherland for assistance
Acknowledgements generating several of the distribution maps.
Lastly, we would like to thank reviewers and
We thank the many curators and collections staff
editorial staff that helped improve the manuscript
that provided loan material or hosted museum
at each stage.
visits facilitating access to vital specimens:
Maxwell Barclay (NHM), Craig Brabant (WIRC),
Michael Brattain, Yves Bousquet (CNC), Michael
Caterino (CUAC), Shawn Clark (BYU), Andrew
Cline (CDFA), Anthony Cognato (MSU),

Fig. 4. Line drawings of the labrum of (A) Omosita, (B) Nitidula.

Fig. 5. Dorsal habitus photographs of the North American species of Nitidula. (A) Nitidula bipunctata (B) Nitidula
carnaria (C) Nitidula flavomaculata (D) Nitidula rufipes (E) Nitidula ziczac

Fig. 6. Dorsal habitus photographs of the North American species of Omosita. (Scale bars = 1mm) A) Omosita
discoidea B) Omosita funestra C) Omosita neartica.
References

Abdell, D. H., Wasti, S. S, and G. C. Hartmann. 1982. Saprophagous arthropod fauna associated with
turtle carrion. Appl. Ent. Zool. 17(3): 301–307.
Adair, T. W., and B. C. Kondratieff. 1996. The occurrence of Nitidula flavomaculata (Coleoptera:
Nitidulidae) on a human corpse. Entomol. News. 107(4): 233–236.
Anderson, G. S. and S. L. VanLaerhoven. 1996. Initial studies on insect succession on carrion in
southwestern British Columbia. J. Forensic Sci. 41(4): 617–625.
Audisio, P. 1993. Coleoptera Nitidulidae-Kateretidae. Fauna d'Italia, vol. 32. Edizioni Calderini.
Bologna, Italy.
Brower, A. V. 2006. Problems with DNA barcodes for species delimitation: ‘ten species’ of Astraptes
fulgerator reassessed (Lepidoptera: Hesperiidae). Syst. Biodivers. 4(2): 127–132.
Connell, W. A. 1984. Nearctic Nitidulidae: Synonymy and Additions since Parsons' Revision
(Coleoptera). The Coleopterists Bulletin. 38(2): 160–164.
De Jong, G. D., and W. W. Hoback. 2006. Effect of investigator disturbance in experimental forensic
entomology: succession and community composition. Med. Vet. Entomol. 20(2): 248–258.
Dodge, H. R. 1937. The occurrence of two European nitidulid beetles in Wisconsin. Entomol. News 48:
284.
Downie, N. M., and R. H. Arnett, Jr. 1996. The Beetles of Northeastern North America. Volume 2.
Sandhill Crane Press, Gainesville, Florida.
Dury, C. 1902. A Revised list of the Coleoptera observed near Cincinnati, Ohio, with notes on localities,
bibliographical references, and description of new species. Journal of the Cincinnati Society of Natural
History. 20: 107–196.
Habeck, D. H. 2002. Family 77: Nitidulidae, pp. 311–315. In R. H. Arnett, Jr., M. C. Thomas, P. E.
Skelley, and J. H. Frank (eds.), American Beetles, Vol. 2. CRC Press, Boca Raton FL.
Hoang, D. T., Chernomor, O., Von Haeseler, A., Minh, B. Q., and L.S. Vinh. 2018. UFBoot2:
improving the ultrafast bootstrap approximation. Mol. Biol. Evol. 35(2), 518–522.
Jelínek, J., Carlton, C. E., Cline, A. R., and R. A. B. Leschen. 2010. 10.26. Nitidulidae Latreille, 1802.
In Leschen R. A. B., Beutel R. G., and J. F. Lawrence (eds.) Handbook of zoology, Vol. IV. Arthropoda:
Insecta, Part 38. Coleoptera, beetles. Walter de Gruyter, Berlin, Germany.

Kalyaanamoorthy, S., Minh, B. Q., Wong, T. K., von Haeseler, A., and L.S. Jermiin. 2017.
ModelFinder: fast model selection for accurate phylogenetic estimates. Nat. Methods. 14(6): 587.
Katoh, K. and D. M. Standley. 2013. MAFFT multiple sequence alignment software version 7:
improvements in performance and usability. Mol. Biol. Evol. 30(4): 772–780.
Kearse M., Moir R., Wilson A., Stones-Havas S., Cheung M., Sturrock S., Buxton S., Cooper A.,
Markowitz S., Duran C., Thierer T., Ashton B., Meintjes P., and A. Drummond. 2012. Geneious
basic: an integrated and extendable desktop software platform for the organization and analysis of
sequence data. Bioinformatics. 28: 1647–1649.
Kirejtshuk 1987. New taxa of the Nitidulidae (Coleoptera) of the East Hemisphere (part 1). Omosita
nearctica sp. n., vicariant with Palaearctic O. colon L. Proc. Zool. Inst. USSR Ac. Sci. 164, 63–94 (in
Russian).
Lord, W. D., and J. F. Burger. 1984. Arthropods associated with herring gull (Larus argentatus) and
great black–backed gull (Larus marinus) carrion on islands in the Gulf of Maine. Environ. Entomol.
13(5): 1261–1268.
Majka, C. G., and A. R. Cline. 2006. Nitidulidae and Kateretidae (Coleoptera: Cucujoidea) of the
Maritime provinces of Canada. I. New records from Nova Scotia and Prince Edward Island. The Canadian
Entomologist. 138(3): 314–332.
McLeod, E. V. H. 2015. The effect of rainfall on blowfly (Calliphoridae) activity and decomposition on
recently deposited animal remains. Doctoral dissertation, Boston University, Boston.
Monthei, D. R. 2009. Entomotoxicological and thermal factors affecting the development of forensically
important flies. Doctoral dissertation, Virginia Tech, Blacksburg.
Nguyen, L. T., Schmidt, H. A., Von Haeseler, A., and B. Q. Minh. 2015. IQ-TREE: a fast and effective
stochastic algorithm for estimating maximum-likelihood phylogenies. Mol. Biol. Evol. 32(1): 268–274.
Parsons, C. T. 1943. A revision of the Nearctic Nitidulidae (Coleoptera). Bulletin of the Museum of
Comparative Zoology 92: 121–278.
Powell, G. S. 2015. A checklist of the sap beetle (Coleoptera: Nitidulidae) fauna of Indiana, with notes on
effective trapping methods. Insecta Mundi. 424: 1–9.
Powell, G. S., Cline, A. R., Duffy, A. G., and J. M. Zaspel. 2020. Phylogeny and reclassification of
Carpophilinae (Coleoptera: Nitidulidae), with insights into the origins of anthophily. Zool. J. Linn. Soc.
189: 1359–1369
Price, M. B. and D. K. Young. 2006. An annotated checklist of Wisconsin sap and short-winged flower
beetles (Coleoptera: Nitidulidae, Kateretidae). Insecta Mundi. 109: 69–84.
QGIS Development Team. 2020. QGIS Geographic Information System. Open Source Geospatial
Project.
Rambaut, A., and A. Drummond. 2008. FigTree: Tree figure drawing tool, version 1.2.2. Institute of
Evolutionary Biology, University of Edinburgh, Edinburgh, UK.
Reed H. B. 1958. A study of dog carcass communities in Tennessee with special reference to the insects.
Am. Midl. Nat. 59: 213–45.
Rodriguez III, W. C. 1982. Insect activity and its relationship to decay rates of human cadavers in East
Tennessee. M.A. Thesis University of Tennessee, Knoxville.
Rubinoff, D., Cameron, S., and K. Will. 2006. A genomic perspective on the shortcomings of
mitochondrial DNA for “barcoding” identification. J. Hered. 97(6): 581–594.
Vogt, G. B. 1950. Occurrence and records of Nitidulidae. The Coleopterists Bulletin 4: 81–91.
Wang, Y., Wang, M., Hu, G., Xu, W., Wang, Y., and J. Wang. 2020. Temperature–dependent
development of Omosita colon at constant temperature and its implication for PMImin estimation. J.
Forensic and Leg. Med. 72:101946.
Watson, E. J., and C. E. Carlton. 2003. Spring succession of necrophilous insects on wildlife carcasses
in Louisiana. J. Med. Entomol. 40(3): 338–347.
Watson, E. J., and C. E. Carlton. 2005. Insect succession and decomposition of wildlife carcasses
during fall and winter in Louisiana. J. Med. Entomol. 42(2): 193–203.
Williams, R. N., M. S. Ellis, and S. Teraguchi. 1997. Nitidulidae (Coleoptera) survey in northeast Ohio.
J. Entomol. Sci. 32: 204–211.
Zanetti, N. I., Visciarelli, E. C., and N. D. Centeno. 2013. Preliminary data on larval morphology and
life cycle of Nitidula carnaria (Coleoptera: Nitidulidae), a species of forensic interest. Revista de la
Sociedad Entomológica Argentina. 72: 3–4.