Biodiversity Data Journal 9: e65768 (eo @)
doi: 10.3897/BDJ.9.e65768 open access

Research Article

Ants of the Palouse Prairie: diversity and species

composition in an endangered grassland

Kayla A Dilwortht, Marek L Borowiec$, Abigail L Cohen, Gabrielle S Mickelsont, Elisabeth C Oeller*,
David W Crowder?, Robert E Clark+

$+ Washington State University, Pullman, United States of America
§ University of Idaho, Moscow, United States of America

Corresponding author: Robert E Clark (robert.e.clark@wsu.edu)

Academic editor: Brian Lee Fisher

Received: 10 Mar 2021 | Accepted: 16 Apr 2021 | Published: 10 May 2021

Citation: Dilworth KA, Borowiec ML, Cohen AL, Mickelson GS, Oeller EC, Crowder DW, Clark RE (2021) Ants of
the Palouse Prairie: diversity and species composition in an endangered grassland. Biodiversity Data Journal 9:
e65768. https://doi.org/10.3897/BDJ.9.e65768

Abstract

Grasslands are globally imperilled ecosystems due to widespread conversion to agriculture
and there is a concerted effort to catalogue arthropod diversity in grasslands to guide
conservation decisions. The Palouse Prairie is one such endangered grassland; a mid-
elevation habitat found in Washington and Idaho, United States. Ants (Formicidae) are
useful indicators of biodiversity and historical ecological disturbance, but there has been no
structured sampling of ants in the Palouse Prairie. To fill this gap, we employed a rapid
inventory sampling approach using pitfall traps to capture peak ant activity in five habitat
fragments. We complemented our survey with a systemic review of field studies for the ant
species found in Palouse Prairie. Our field inventory yielded 17 ant species across 10
genera and our models estimate the total ant species pool to be 27. The highest ant
diversity was found in an actively-managed ecological trust in Latah County, Idaho,
suggesting that restoration efforts may increase biodiversity. We also report two rarely-
collected ants in the Pacific Northwest and a microgyne that may represent an undescribed
species related to Brachymyrmex depilis. Our score-counting review revealed that
grassland ants in Palouse Prairie have rarely been studied previously and that more ant
surveys in temperate grasslands have lagged behind sampling efforts of other global
biomes.

© Dilworth K et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY
4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are
credited.

2 Dilworth K et al

Keywords

Palouse Prairie, ant biodiversity, rapid species inventory, agroecosystems, insect
conservation, prairies

Introduction

Temperate grasslands and savannah are amongst the most endangered biomes in the
world, having the highest rate of conversion to agriculture and the lowest rate of
government protection (Hoekstra et al. 2005). In the United States, grasslands, savannah
and barrens communities are critically endangered, experiencing > 98% decline in areas
since European settlement (Noss and Scott 1995). Since organisms found in prairies are
threatened by habitat loss, biodiversity surveys that describe the native and non-native
fauna are necessary to help provide information for future conservation efforts and guide
restoration to areas subject to impacts. Ants are useful biodiversity indicators in grasslands
and characterising ant community composition through rapid assessment is pertinent for
monitoring and evaluating global grassland restoration efforts (Peters et al. 2016).

Biodiversity surveys play a strategic role in grassland conservation, aiding arguments that
habitats contain rare or endemic species (Van Schalkwyk et al. 2019). Temperate
grasslands are host to a wide range of endemic plant and animal communities, including
many insects of conservation concern (Sampson and Knopf 1994). A comprehensive list of
globally-threatened species, the Red List, counts over 2,000 grassland species that are
critically engendered, endangered or vulnerable worldwide, 108 of which are in North
America (I.U.C.N. 2020). Grasslands are frequently fragmented by agricultural production
and they are, therefore, prone to increased rates of local extinctions through a variety of
modes, such as reduced population sizes, increased invaders and elimination of keystone
predators (Leach and Givnish 1996).

The Palouse Prairie is an endangered grassland ecosystem that originally encompassed
south-eastern Washington State and neighbouring northern Idaho (Looney and Eigenbrode
2012, Noss and Scott 1995). The Palouse landscape is characterised by rolling hills
formed from wind-blown fertile loess soils, which form the foundation for plant communities
of caespitose grass, co-dominant shrubs and forbs (Looney et al. 2009, Daubenmire
1970). The Palouse Prairie has experienced an estimated 99.9% decline in habitat across
its former range due to agricultural land use, making it one of the most imperilled
ecosystems in the United States (Donovan et al. 2009, Black et al. 1998). What habitat
remains is fragmented into small, narrow strips of land and a majority of these fragments
are smaller than two hectares with high perimeter-area ratios (Yates et al. 2004).
Encouragingly, these fragments are capable of supporting several endangered species
(Looney and Eigenbrode 2012).

Published biodiversity surveys are limited for the Palouse Prairie, encompassing a survey
of bumblebees (Hatten et al. 2013), macromoths (Thompson et al. 2014) and forbs
(Donovan et al. 2009). In this study, we chose ants (Formicidae) for the biodiversity survey

Ants of the Palouse Prairie: diversity and species composition in an endangered ... 3

in Palouse Prairie. Ants are used prominently for monitoring because they are ubiquitous in
terrestrial ecosystems, sampling is relatively inexpensive and many ant species respond
quickly to disturbances (Tiede et al. 2017, Hoffmann 2010, Underwood and Fisher 2006,
Folgarait 1998, Andersen 1990). However, in poorly-surveyed regions, using ants as
biodiversity indicators can be challenging if taxonomic resources are not well developed
(Hevia et al. 2016). The Pacific Northwest Region is generally under-sampled for ants and
thus up-to-date taxonomic resources are limited (Hoey-Chamberlain et al. 2010).
Consequently, we included a systematic review of all ant species found and referred
directly with taxonomic experts to verify all species-level identifications.

Material and methods
Rapid inventory survey

Ant surveys were completed at five prairie fragments: Hudson Biological Reserve, Kamiak
Butte County Park, Skinner Ecological Preserve, Idler’s Rest Nature Preserve and Philips
Farm County Park (Figs 1, 2, Suppl. material 1). We used a rapid inventory sampling
method with pitfall trap transects amongst the five sites (Agosti et al. 2000, Ellison et al.
2007). We used 4 cm diameter, 14 cm depth, pitfall traps filled 2/3 with propylene glycol
(Higgins and Lindgren 2012). Our survey intended to capture peak ant activity in August on
days above 24°C with no precipitation. We worked in four teams to ensure all traps were
placed and collected simultaneously. Pitfall traps were run for 48 h, after which ants were
transferred to 95% ethanol. Afterwards, ants were pinned for later identification and
storage. In total, we ran 131 pitfall traps and collected 424 individual ants (Suppl. material
2).

Figure 1. EES]

Example of an intact Palouse Prairie habitat fragment located at the Hudson Biological
Preserve (“Smoot Hill”) in Albion, Washington.

4 Dilworth K et al

Species-level identification of ants can be time-consuming and difficult in regions with
poorly-described ant fauna (reviewed in Ellison 2012). The inland North-western US has
few recent records of ants to aid in identification and we thus employed a four-step
identification approach. First, all ants were identified to genus using Ant Genera of North
America (Fisher and Cover 2007). Second, we used records and pinned specimen
photographs posted to online ant taxonomy databases (i.e. Guenard et al. 2017) to identify
ants to species. Third, we compared specimens to collections at the Washington State
University MT James Museum and the University of Idaho WF Barr Entomological
Museum. We also consulted published checklists of ants known from Idaho, Washington
and the western United States (Cole 1936, Smith 1941, Yensen et al. 1977, Cook 1953,
Wheeler and Wheeler 1986), although these studies reflect the lack of recent
comprehensive reports published.

46.9 Kamiak Butte

Phillips-Fatm~™

46.6 aa — <<
__ -Idler's Rest

==

es

Latitude

46.7 Seas, eee
bg Cees a
7

”_. Skinner Preserve = =

46 ; —
717.3 -117.2 -117.1 -117.0 -116.9 -116.8
Longitude

Figure 2. EES

Map of survey locations for the five Palouse Prairie sites. Red circles encompass the prairie
fragment sampled. Inset shows map extent in the Pacific Northwest.

Systematic review methods

We searched for studies in Web of Science that quantified abundance or richness of the
ant species within prairie or grassland from 2016 to the present. Our search was
conducted in June 2020 using the term “ant AND divers* AND prairie OR ant AND divers*
AND grassland”. Our search yielded 106 studies that were reviewed for inclusion, based
on three criteria: (i) the study assessed more than one species; (ii) the study was
performed in non-agricultural grassland or prairie; and (iii) the biome of the study could be
determined; 21 studies met these criteria (Suppl. material 3). We classified the biome of
each study using either the site coordinates or other geographic descriptors to locate study
areas, then cross-referenced with the EcoRegions web app (Dinerstein et al. 2017).

We examined the frequency of recent publications on each individual species collected in
our pitfall survey using the same approach as the biome survey, but instead used the
genus and species names as search terms. We tabulated the number of studies from

Ants of the Palouse Prairie: diversity and species composition in an endangered ... 5

2010-2020 that reported ant species we found. We read abstracts to ensure that all studies
included in this survey involved field observations of the insect species in question and we
included multiple species names for recently-revised species (Schar et al. 2018). We did
not include purely lab-based studies on behaviour (i.e. Bordoni et al. 2019).

Statistical methods

Analyses and figure generation were completed in R ver 4.0.2 (CRAN 2020). Visualisation
of sites was mapped using ggpmap package (Kahle and Wickham 2013). Species
accumulation curves were run, using the ‘vegan’ package in R (Dixon 2003). We estimated
species richness using the Chao1 estimate, an abundance-based estimate of species
richness, following decision-tree recommendations (Hortal et al. 2006).

Results

Amongst all sites, we collected 17 ant species (Table 1), with Aohaenogaster occidentalis
the most common. Ants in the genus Formica were the most diverse, with six species (Fig.
3). The non-native species Tetramorium immigrans was found in three locations, but was
not abundant. We found a single microgyne (miniature queen) similar to Brachymyrmex
depilis. Scattered records of similar Brachymyrmex microgynes exist and they have been
interpreted as undescribed, socially parasitic species (Moreau et al. 2014, Deyrup 2016).
Finally, we collected Temnothorax nevadensis and Formica puberula, two ant species that
are rarely collected in Pacific Northwest temperate ecosystems (Gueénard et al. 2017).

Table 1.
Studies reporting species found in this survey between, published 2010 and 2020.

Ant species Studies Paper References

Aphaenogaster 1 Thomas et al. 2020

occidentalis

Brachymyrmex depilis 3 King and Tschinkel 2016, Rosas-Mejia et al. 2019, Underwood and Christian
2009

Brachymyrmex sp. 2 Moreau et al. 2014, Deyrup 2016

microgyne

Camponotus vicinus 1 Davenport et al. 2013

Formica argentea 2 Lenz et al. 2013, Ouellette et al. 2010

Formica lasioides 1 Karban et al. 2015

Formica neoclara 0

Formica neogagates 5 Clark and Singer 2018, Ellison et al. 2007, Wang et al. 2001, Weseloh 2000,

Weseloh 2001

6 Dilworth K et al

Ant species Studies Paper References

Formica puberula 0

Formica 0

subaenescens

Lasius americanus, 11 Akyirek et al. 2016, Barton and Ives 2014, Cuesta-Segura et al. 2012, Molnar et
reported as Lasius al. 2018, Paluh et al. 2015, Park and Moon 2020, Pech 2014, Warren et al. 2012,
alienus in older works Yitbarek et al. 2011, Zhu and Wang 2018

Prenolepis imparis 13 Abbate and Campbell 2013, Burford et al. 2018, Campbell et al. 2019, Cuautle et

al. 2016, Fitzgerald and Gordon 2012, Gordon and Heller 2014, Kjar and Park
2016, Rowles and Silverman 2009, Sanchez-Pefia 2013, Sorrells et al. 2011,
Taylor et al. 2013, Thomas et al. 2020, Vonshak and Gordon 2015

Solenopsis molesta 4 Guyer et al. 2018, Pecarevic et al. 2010, Resasco et al. 2014, Rojas et al. 2014

Tapinoma sessile 20 Buczkowski and Bennett 2009, Buczkowski 2010, Buczkowski and Krushelnycky
2012, Buczkowski and Richmond 2012, Gibson et al. 2019, Gow et al. 2013,
Hamm 2010, Kaspari et al. 2010, Kautz et al. 2017, Kimball 2016a, Kimball
2016b, Nelson et al. 2020, Neumann and Pinter-Wollman 2019, Ouellette et al.
2010, Powell et al. 2009, Salyer et al. 2014, Schmidt et al. 2012, Scholes and
Suarez 2009, Toennisson et al. 2011, VanWeelden et al. 2015

Temnothorax 1 Adams et al. 2018

nevadensis

Temnothorax 11 Bowens et al. 2013, Blonder and Dornhaus 2011, Cao 2013, Charbonneau et al.
rugatulus 2015, Charbonneau et al. 2017, DiRienzo and Dornhaus 2017, Doering and Pratt

2016, Doering and Pratt 2019, Doering et al. 2020, Doering et al. 2019, Heinze
and Rueppell 2014

Tetramorium 23 Brandt et al. 2010, Cerda et al. 2009, Chin and Bennett 2018, Collignon and
immigrans and Detrain 2010, Cordonnier et al. 2019a, Cordonnier et al. 2020a, Cordonnier et al.
Tetramorium 2020b, Cordonnier et al. 2019b, Dash and Sanchez 2009, Helms et al. 2020,
caespitum Hoey-Chamberlain et al. 2010, Joharchi et al. 2012a, Joharchi et al. 2012b, Li et

al. 2019, Lundgren et al. 2010, Peéarevié et al. 2010, Sheard et al. 2020, Slipinski
et al. 2012, Steiner et al. 2010, Wagner et al. 2017, Wagner et al. 2018a, Wagner
et al. 2018b, Joharchi and Halliday 2013

Ant species richness varied amongst sites, with the highest diversity at Skinner Ecological
Preserve (Fig. 4) and a total predicted species pool of 27 (Chao7 index = 27.9, SE = 10.2).
Skinner Preserve, the largest intact prairie fragment, had the highest species richness (16)
and estimated species pool (Chao7 index = 25.8, SE = 10.0). Smoot Hill had the lowest
species richness (4) and smallest species pool (Chao7 index = 5.9, SE = 3.6). Two
restored habitats adjacent to Ponderosa Pine — Douglas-Fir forests near Moscow, Idaho
had intermediate species pools (Philips Farm [9 species] — Chao7 index = 18.0, SE = 9.2;
Idler’s Rest [13.6 species] — Chao7 index = 13.6; SE = 4.8). Kamiak Butte had 11 species
(Chao7 index = 13.9; SE = 4.4).

Ants of the Palouse Prairie: diversity and species composition in an endangered ... 7

Aphaenogaster occidentalis
Brachymyrmex depilis
Brachymyrmex sp
Camponotus vicinus
Formica argentea

Formica lasioides

Formica neoclara

Formica neogagates
Formica puberula

Formica subaenescens

Lasius americanus*

>
oO
fa
=
n
oO
a
3
2
o
a
=
ce)
=
n
2
S
¢

Prenolepis imparis
Solenopsis molesta
Tapinoma sessile
Temnothorax nevadensis
Temnothorax rugatulus

Tetramorium immigrans**

0) 30 60 90 120
Cumulative number ants collected in pitfalls

Figure 3. EES

Numbers of ants collected in all pitfalls for each species. Asterisks indicates a species that has
recently been revised, thus the literature search includes *Lasius neoniger and ** Tetramonum
caespitum. Brachymyrmex sp. indicates abundance of the Brachyrmex sp. microgyne
collected.

Site

+ @ Idler's Rest

s&h ~Kamiak Butte

® Phillips Farm

ee Skinner Preserve
& Smoot Hill

Ant Species

20 30
# of Pitfalls

Figure 4. EES]

Species accumulation curve with expected mean species richness plotted for each survey
location. Dots and lines show estimated species richness at a given sampling interval, while
shaded area shows 95% Cl.

We found 95 publications detailing a study containing at least one species observed in our
survey (Table 1). No field studies have been published for Formica neoclara, F. puberula
and F. subaenescens and only one study included Temnothorax nevadensis (Table 1).
These limited numbers show our pitfall sampling found several rarely studied or unstudied
ant species. In fact, there were five or fewer publications for all the ant species collected
besides the common “tramp species” or urban pests, like Tapinoma sessile and

8 Dilworth K et al

Tetramorium immigrans (Kamura et al. 2007, Uno et al. 2010). Our systematic review of
ant biodiversity surveys in different biomes over the last five years revealed a bias towards
Tropical & Subtropical Grasslands, Savannahs & Shrublands (eight studies, Cross et al.
2016, Van Schalkwyk et al. 2019, Arcoverde et al. 2016, Lasmar et al. 2020, Drése et al.
2019, Hlongwane et al. 2019, de Queiroz et al. 2020, Santoandré et al. 2019). We found
two studies on Tropical & Subtropical Moist Broadleaf Forests (Lawes et al. 2017, Klunk et
al. 2018), one study on Tropical & Subtropical Coniferous Forests (Cuautle et al. 2016),
three studies on Temperate Broadleaf & Mixed Forests (Braschler and Baur 2016, Helms
et al. 2020, Heuss et al. 2019), one study on Montaine Grasslands & Shrublands (Jamison
et al. 2016), three studies on Mediterranean Forests, Woodlands & Scrub (Adams et al.
2018, Catarinue et al. 2018, Flores et al. 2018) and one study on Deserts & Xeric
Shrublands (Alvarez and Ojeda 2019). In the last five years, only two citations have
included information on ants in Temperate Grasslands, Savannahs and Grassland biomes
(Ramos et al. 2018, Kim et al. 2018), the biome that encompasses Palouse Prairie (Fig. 5).

Tropical & Subtropical Moist Broadleaf Forests
Tropical & Subtropical Grasslands, Savannas & Shrublands
Tropical & Subtropical Coniferous Forests

Temperate Grasslands, Savannas & Shrublands

Biomes

Temperate Broadleaf & Mixed Forest

Montane Grasslands & Shrublands

Mediterranean Forests, Woodlands & Scrub

Deserts & Xeric Shrublands

7.5 10.0

2.5 5.0
Number of publications since 2015

Figure 5. EES

Number of ant biodiversity surveys published over a 5-year period from 2016-2020 where
locations were reported. Biome classifications inferred from a global terrestrial ecoregion map
(Dinerstein et al. 2017).

Discussion

Our model results suggest there are likely many more ant species to be discovered with
more intensive sampling in Palouse Prairie and that there is likely to be appreciable
diversity of other insect species beyond those of the limited taxonomic focus of this study.
Furthermore, one of our collected specimens may be a currently undescribed species of
ant, but this putative ant species is rarely collected and taxonomic revision of
Brachymyrmex is required to demonstrate if this is the case. In many ecosystems, insect
faunas are poorly described (Berenbaum 2008, Dunn 2005). A lack of recent publications
on temperate grassland ant faunas and dearth of work on several species collected in

Ants of the Palouse Prairie: diversity and species composition in an endangered ... ss)

Palouse Prairie underscore the importance of survey work in endangered ecosystems. As
habitat destruction and fragmentation continue, we may never be able to sample or study
the more rarely-collected species (Dunn 2005, Noss and Scott 1995).

Ant communities are under-sampled in cool-temperate ecosystems compared to the
tropics and sub-tropics, including temperate grasslands in the north-western United States
(Ellison 2012, Radtke et al. 2014). Since ants are excellent biological indicators of
ecosystem health, sampling efforts may use our data as a comparison point to see if
restoration efforts have been successful (Folgarait 1998, Williams 1994). Luckily, once
taxonomic resources are available, assessment of an ecosystem’s ant communities can be
completed quickly with greater accuracy of species-level identification (Ellison et al. 2007).
The intermediate levels of ant diversity at sites adjacent to forest validate predictions that
Palouse Prairie-forest ecotones may support high biodiversity (Morgan et al. 2020) Finally,
our estimates of a species pool of 27 reflect similar scales of ant species richness found in
large ant surveys in grassland systems, such as Wisconsin, USA tallgrass prairie (29
species in control sites, Kim et al. 2018) and Argentinian grasslands (46 species in
grassland sites, Santoandre et al. 2019). However, given our absolute species richness
was 17, there is more sampling to be completed to comprehensively describe this fauna. In
fact, follow up hand-collect events at Skinner Preserve completed in 2019 found five
additional ant species, including Formica altipetens, Formica aserva, Formica obscuripes,
Formica ravida and Lasius interjectus (Borowiec, unpublished data).

Research in conservation biological control has shown the value of ants as predators in
agroecosystems (Way and Khoo 1992). In our study, we found multiple species of ants in
genera often implicated as predators of chewing herbivores, such as Formica and
Camponotus (Drummond and Choate 2011). More recent work demonstrates that social
insects, including ground-nesting ants, have positive effects on dryland crop yield by
increasing water and micronutrient availability to cereals (Evans et al. 2011). However, the
value of these ecosystem services likely pales in comparison to the benefit ants could
provide as predators of weed seeds (e.g. Evans and Gleeson 2016). Weeds and herbicide
resistance are amongst the most economically-challenging pest problems in dryland
agriculture (Powell and Shaner 2001) and ants have been implicated in regulating seed
banks in other grasslands (Motze et al. 2013). Recovery of more Palouse Prairie could
promote higher ant diversity and abundance (Lawes et al. 2017), thus increasing the
likelinood that ants are available to provide these critical ecosystem services, including
weed seed predation.

Conclusions

The remaining habitat in the Palouse is highly fragmented and only protected by a range of
public and private trusts (Looney and Eigenbrode 2012). This is problematic since we
found several species of ants that are uncommonly collected. While these species are not
endemic to Palouse Prairie, our observations suggest that opportunities to study these
insects in grassland habitats are limited. Furthermore, in addition to fragmentation and
disturbance from agriculture, several of our sites are invaded by the pavement ant (

10 Dilworth K et al

Tetramorium immigrans), which is associated with disturbed, agricultural habitats (Cerda et
al. 2009). The extremely reduced range of the Palouse Prairie and the presence of
invasive species means many other taxonomic groups of animals are in urgent need of
sampling before opportunities to describe this fauna are irreversibly lost.

Author contributions

KAD, ALC, ECO and REC completed field surveys. REC analysed data. KAD, ALC, GSM
and ECO performed the systematic review. KDA, REC and MLB curated and identified
ants. All authors contributed to writing and editing the manuscript.

Conflicts of interest

The authors declare no conflicts of interest.

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Ants of the Palouse Prairie: diversity and species composition in an endangered ...

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