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The ants of Ohio (Hymenoptera, Formicidae): an updated checklist
expand article infoKaloyan Ivanov
‡ Virginia Museum of Natural History, Martinsville, United States of America
Open Access

Abstract

I update the last published list of Ohio ants to include 26 new species records and 38 name changes in species already on the list based on literature records, institutional collections, and contemporary collections made by the author or colleagues. At present, 143 species and morphospecies representing 30 native and 5 exotic genera and 7 subfamilies have been recorded for the state. Another seven species are removed from the Ohio ant fauna as they represent distribution anomalies, or are based on erroneous records. Known distribution data suggest that there is still a considerable potential for the discovery of more ant taxa in Ohio.

Keywords

ant diversity, new records, distribution, eastern United States, Nearctic

Introduction

In 2005, Coovert published the first comprehensive account of the ant fauna of Ohio and listed 118 species and subspecies from the state based on literature, museum records, and extensive collecting in the state in the period 1996–2002 (Coovert 2005). The list included 14 taxa recorded for the first time from Ohio. A few earlier records were overlooked (i.e., Paratrechina longicornis (Latreille) (Hedges 1998), Strumigenys hyalina (Bolton) (Bolton 2000), Tetramorium bicarinatum (Nylander) (Bolton 1979)), or excluded from the list (i.e., Formica incerta Buren (Wesson and Wesson 1940; Amstutz 1943; Headley 1943, 1949, 1952; Talbot 1945), Myrmica incompleta Provancher (Weber 1950), Proceratium crassicorne Emery (Wesson and Wesson 1940)) based on contemporary understanding of the group. Formica nitidiventris Emery was later synonymized with Formica pallidefulva Latreille and dropped from Ohio’s list (Trager et al. 2007), bringing the total number of species known from the state to 117.

In the past fourteen years, a substantial amount of information from survey work, biodiversity and ecological studies, and taxonomic revisions (Francoeur 2007; Rabeling et al. 2007; Snelling and Snelling 2007; Trager et al. 2007; Trager 2013; Mackay and Mackay 2010, 2017; Kallal and Lapolla 2012; Pacheco and Mackay 2013; DeMarco and Cognato 2015, 2016; Morgan and Mackay 2017) has become available. As a result, the 2005 list is much out of date necessitating the current update. Contemporary knowledge of the state’s ant fauna may be beneficial in providing support to future ecological, conservation, behavioral, and taxonomic studies in Ohio and the surrounding areas.

Materials and methods

Data reported herein are based on information gathered from the following sources: 1) review of published records; 2) examination of materials in the collections of the Cleveland Museum of Natural History and the Virginia Museum of Natural History; 3) online records from digitized personal and museum collections, iDigBio (https://www.idigbio.org/), SCAN (http://scanbugs.org/portal), and AntWeb (https://antweb.org/); and 4) newly collected material.

Abbreviations of entomological collections used in this study are as follows:

CMNH Cleveland Museum of Natural History, Cleveland, Ohio.

FMNH Field Museum of Natural History, Chicago, Illinois.

INHS Illinois Natural History Survey, Champaign, Illinois.

MCZ Louis Agassiz Museum of Comparative Zoology, Cambridge, Massachusetts.

MEM Mississippi Entomological Museum, Starkville, Mississippi.

OSUC C.A. Triplehorn Insect Collection at Ohio State University, Columbus, Ohio.

TTU-Z Texas Tech University Invertebrate Collection, Lubbock, Texas.

UAIC University of Arizona Insect Collection, Tucson, Arizona.

UCMC University of Colorado Museum of Natural History Insect Collection, Boulder, Colorado.

VMNH Virginia Museum of Natural History, Martinsville, Virginia.

To the best of my knowledge, all species names presented herein are the currently valid names according to the fourth edition of the International Code of Zoological Nomenclature (ICZN 1999). Taxon names and concepts used in this work are gleaned from an interpretation of the contemporary literature and follow Bolton (2019) with the exception of Myrmica emeryana Forel which is based on an ongoing revision of North American Myrmica by André Francoeur. Known morphospecies bear the names assigned to them by the relevant taxonomic specialists (e.g., André Francoeur’s “Myrmica AF-smi”). These names are unavailable according to the zoological code (ICZN 1999). Unless otherwise noted, all identifications were made by the author. Vouchers of all newly collected material are deposited in the invertebrate collections at CMNH and VMNH.

The species included in this checklist can be identified using the keys in Coovert (2005), Francoeur (2007), Trager et al. (2007), Ellison et al. (2012), Kallal and Lapolla (2012), and Sarnat et al. (2015). Additionally, representative high-resolution photographs of at least the worker’s caste of Ohio’s species can be found on AntWeb, AntWiki (www.antwiki.org) and on Discover Life (www.discoverlife.org).

In the species accounts presented herein, all taxa are listed alphabetically by subfamily, tribe, genus, and species. County-level distributions (new data in bold) are provided only for the published Ohio records not appearing in Coovert (2005) and for species previously known from only one or two records which are herein presented as “Rare”, although I acknowledge that they may be more abundant and widespread outside the boundaries of Ohio. Detailed locality data are given only for the unpublished Ohio records. Material from institutional and personal collections is given first, followed by literature records, which are presented in chronological order. An asterisk (*) marks taxa absent from the 2005 list and superscript E denotes exotic species. General within-state distribution follows Coovert (2005) with adjustments for new records and recent taxonomic changes.

Appended is a list of questionable Ohio records based on published accounts or unpublished records appearing in online databases. These taxa are excluded from the Ohio fauna as they are based on erroneous records, mislabeled specimens, or misidentifications.

Results and discussion

A total of 143 species and morphospecies representing 30 native and 5 exotic genera and 7 subfamilies comprise the known ant fauna of Ohio, USA. These include 26 additions to the checklist provided in Coovert (2005) and 38 name changes in species already on the list. The new Ohio records are based on: 1) records published prior to 2005 but previously omitted from the list (6 taxa), 2) records published since 2005 (10 taxa), and 3) unpublished records (10 taxa). The majority of Ohio’s new additions are known species whose populations were only recently discovered in the state with the remainder representing recently described (Myrmica semiparasitica Francoeur) or yet undescribed (Myrmica AF-eva, Myrmica AF-scu, Myrmica AF-smi) species.

Ohio’s ant fauna is dominated by cool-climate Camponotus, Formica, Lasius, and Myrmica, whereas taxa with austral affinities are largely restricted to the southern, unglaciated region of the state. The overall composition of Ohio’s myrmecofauna much resembles that of other Midwestern and northeastern states for which contemporary checklists exist (e.g., Indiana (Carroll 2011), Michigan (Wheeler et al. 1994), New England (Ellison et al. 2012), and Pennsylvania (Butler and Coulter 2019)). The most species-rich genera in the state include Formica (26 spp.), Strumigenys (15 spp.), Myrmica (11 spp. and morphospp.), Camponotus (10 spp.), Lasius (10 spp.), Temnothorax (10 spp.), Aphaenogaster (7 spp.), and Nylanderia (6 spp.). These eight genera include two-thirds of Ohio’s ant species and morphospecies. Conversely, 16 genera are represented in Ohio by a single species (Table 1).

Taxonomic composition of Ohio’s ant fauna. Genera are ordered by the total number of (morpho-)species in each. Superscript E denotes exotic genera.

Genus Number of species % of Ohio fauna Genus Number of species % of Ohio fauna
Formica 26 18.2 Tapinoma 2 1.4
Strumigenys 15 10.5 Brachymyrmex 1 0.7
Myrmica 11 7.7 EBrachyponera 1 0.7
Camponotus 10 7.0 ECardiocondyla 1 0.7
Lasius 10 7.0 Dorymyrmex 1 0.7
Temnothorax 10 7.0 Forelius 1 0.7
Aphaenogaster 7 4.9 Harpagoxenus 1 0.7
Nylanderia 6 4.2 Leptothorax 1 0.7
Pheidole 5 3.5 ELinepithema 1 0.7
Stenamma 4 2.8 Myrmecina 1 0.7
Crematogaster 3 2.1 Neivamyrmex 1 0.7
Dolichoderus 3 2.1 EParatrechina 1 0.7
Proceratium 3 2.1 Polyergus 1 0.7
Solenopsis 3 2.1 Ponera 1 0.7
ETetramorium 3 2.1 Prenolepis 1 0.7
Colobopsis 2 1.4 Stigmatomma 1 0.7
Hypoponera 2 1.4 Trachymyrmex 1 0.7
Monomorium 2 1.4

Many of Ohio’s ant taxa are widespread, both in the state and in Eastern North America, with nearly a quarter of the species having an essentially statewide distribution. The most widely distributed of these are the forest-dwelling Camponotus pennsylvanicus (De Geer) and Lasius americanus Emery, which have been recorded in all of Ohio’s 88 counties. Other species that have been collected in a large proportion of the state include Lasius neoniger Emery (87 counties), Formica subsericea Say (>80 counties), Formica pallidefulva Latreille (>75 counties), and the non-native Tetramorium immigrans Santschi (>75 counties), (distribution from Coovert 2005 and new data). Other taxa exhibit restricted within-state distribution as a result of associations with rare/unusual habitats (Dorymyrmex grandulus (Forel), Nylanderia arenivaga Wheeler, W.M., Myrmica lobifrons Pergande), or because they reach the northern (e.g., Neivamyrmex carolinensis (Emery), Aphaenogaster lamellidens Mayr, Crematogaster pilosa Emery, Pheidole tysoni Forel, Strumigenys laevinasis Smith, M.R., Temnothorax pergandei (Emery), Trachymyrmex septentrionalis (McCook)), or southern (e.g., Formica aserva Forel, Formica ulkei Emery, Harpagoxenus canadensis Smith, M.R., Leptothorax canadensis Provancher) limits of their distributional ranges. Yet others, have been infrequently collected in the state due to their socially parasitic lifestyle (F. aserva, Temnothorax americanus (Emery), T. duloticus (Wesson L.G.), T. minutissimus Smith M.R., Tetramorium atratulum (Schenck)), or simply because standard ant collecting techniques do not pick them very well (e.g., Colobopsis, Strumigenys, Proceratium). Undoubtedly, there are other, equally rare, species, which have not been detected yet in Ohio.

Recently, Ivanov (2016) reviewed the exotic ant fauna of Ohio and listed ten non-native taxa from the state. Earlier Ohio records of Tetramorium bicarinatum (Bolton 1979; Wetterer 2009) were overlooked and not included in the list. With the addition of this species, and the recent report of Brachyponera chinensis (Emery) from Cincinnati (Guénard et al. 2018), there are now 12 ant exotics known to occur in the state. Half of Ohio’s non-native species belong to the subfamily Myrmicinae (6 species) with Dolichoderinae, Formicinae, and Ponerinae represented by two species each. While the native ant fauna predominates in intact forest ecosystems, grasslands, old fields, and prairie remnants, exotic species are more commonly associated with disturbed areas and are frequently encountered indoors in the large metropolitan areas of Cincinnati, Cleveland, and Columbus.

Although the updated list presented herein likely contains a large proportion of the taxa that exist in the state, there is still a considerable potential for the discovery of more species. There are at least 44 species likely to occur in the state by virtue of their presence in neighboring areas (Appendix 1), any and all of which may turn up in Ohio in the future. Targeting these taxa, however, will require a more systematic approach and more intensive collecting throughout the state. Nonetheless, future collecting efforts are certain to add new taxa and provide additional locality records of known species building upon our knowledge of Ohio’s myrmecofauna.

List of Ohio ants

Subfamily Amblyoponinae [1 genus; 1 species]

Tribe Amblyoponini

Stigmatomma Roger

pallipes (Haldeman). Common, but inconspicuous element of the local fauna throughout Ohio. Under Amblyopone in Coovert (2005).

Subfamily Dolichoderinae [5 genera; 8 species]

Tribe Dolichoderini

Dolichoderus Lund

plagiatus (Mayr). Widespread.

pustulatus Mayr. Widespread

taschenbergi (Mayr). Champaign and Hocking Cos. in southcentral Ohio. Rare.

Tribe Leptomyrmecini

Dorymyrmex Mayr

grandulus (Forel). Literature records: Lucas (Friedrich 2010). Only known from a small number of collections in prairie remnants of the globally rare Oak Openings Region of northwestern Ohio. Rare.

Forelius Emery

pruinosus (Roger). Literature records: Lucas, Preble (Uno et al. 2010; Campbell and Crist 2017). Previously reported from areas with full sun and hot, dry conditions in Adams and Erie Cos. Rare.

Linepithema Mayr

Ehumile (Mayr). Ohio’s only confirmed record of this notorious invasive species is from a greenhouse in Cleveland (Ivanov 2016). Coovert (2005) includes an unspecified Ohio record in Arnett (1993). Indoors, in heated buildings.

Tribe Tapinomini

Tapinoma Foerster

*, Emelanocephalum (Fabricius). Literature records: Butler, Cuyahoga, Franklin (Ivanov 2016). In Ohio, this widely distributed tramp species is currently confined to indoor situations and can be abundant in greenhouses, conservatories, and zoo buildings.

sessile (Say). Statewide and in most habitats including disturbed sites and inside buildings.

Subfamily Dorylinae [1 genus; 1 species]

Neivamyrmex Borgmeier

carolinensis (Emery). Unspecified Ohio records in M.R. Smith (1967), Watkins (1972, 1976), and D.R. Smith (1979). Rare. No definitive Ohio records of this southeastern army ant exist, but it may occur in suitable habitats in the southern part of the state along the northern fringes of its distribution (see Snelling and Snelling 2007).

Subfamily Formicinae [9 genera; 58 species]

Tribe Camponotini

Camponotus Mayr

americanus Mayr. Widespread. Majority of the records from unglaciated southern Ohio, with disjunct records from Columbiana, Lucas (Uno et al. 2010) and Summit (VMNH110593) Cos. in the north.

caryae (Fitch). Widespread.

castaneus (Latreille). A number of scattered records from southern Ohio, north to Franklin and Muskingum Cos.

chromaiodes Bolton. Statewide.

discolor (Buckley). Gallia and Muskingum Cos. in unglaciated southwestern Ohio. Rare.

*herculeanus (Linnaeus). – Cuyahoga Co. • Cuyahoga Valley National Park; 41.35597N, 81.56847W; alt. 259 m; 18 July 2009; W. & E. Mackay leg.; #23734; nest in solid log; hardwood forest; clay loam soil; W. Mackay det. 2009; MCZENT567833.

nearcticus Emery. Statewide.

novaeboracensis (Fitch). Widespread in northern Ohio, south to Franklin and Greene Cos.

pennsylvanicus (De Geer). Statewide.

subbarbatus Emery. Statewide.

Colobopsis Mayr

impressa Roger. Mahoning (CMNHENT39502), Warren (K. Campbell unpublished data; pers. comm., February 2019). Previously reported from Henry and Muskingum Cos. Rare.

mississippiensis (Smith, M.R.). This infrequently collected arboreal species is known from only a few scattered Ohio sites.

Tribe Formicini

Formica Linnaeus

argentea Wheeler, W.M. Cuyahoga, Ottawa, Richland (MCZENT551348, MCZENT551349, MCZENT551350, MCZENT551351, MCZENT551352, MCZENT551353, MCZENT551354, MCZENT551355, MCZENT551400, MCZENT551401, MCZENT551402, MCZENT551410, MCZENT551411, MCZENT551412, MCZENT551413). Until now, this species was only know Summit Co. in the northeastern part of the state. Rare.

aserva Forel. Geauga (VMNH110555; in a mixed colony with host F. glacialis Wheeler, W.M). Until present, this northern dulotic species was only known from neighboring Ashtabula Co. in the extreme northeastern part of the state. Rare.

dakotensis Emery. Scattered records from glaciated western Ohio.

difficilis Emery. Single record from Jackson Co. in southcentral Ohio (Wesson and Wesson 1940). Rare.

dolosa Buren. Scattered sites throughout Ohio. Records under Formica schaufussi Mayr in Coovert (2005) refer to this species.

exsectoides Forel. Statewide.

glacialis Wheeler, W.M. Widespread in glaciated northern Ohio, south to Champaign and Madison Cos.

*gynocrates Snelling, R.R. & Buren. Literature records: Lucas (Friedrich 2010). In Ohio, this species is only known from prairie remnants in the globally rare Oak Openings Region of northwestern Ohio.

*incerta Buren. Ashland, Ashtabula, Athens, Erie, Geauga, Hardin, Huron, Madison, Morgan, Summit (CMNHENT40013, CMNHENT40014, CMNHENT40015, CMNHENT40016, CMNHENT40017, CMNHENT40018, CMNHENT40019, CMNHENT40259, MCZENT672736, MCZENT672737, VMNH110556, VMNH110557, VMNH110558, VMNH110575, VMNH110576, VMNH110597, VMNH110601, VMNH110602). Literature records: Adams, Ashtabula, Butler, Champaign, Delaware, Hamilton, Hocking, Jackson, Lucas, Montgomery, Seneca, Wyandot (Amstutz 1943; Headley 1943, 1949, 1952; Talbot 1945; Trager et al. 2007; Friedrich 2010), southcentral Ohio (Wesson and Wesson 1940). The name incerta has a long and confusing history (see Coovert 2005; Trager et al. 2007) until finally revived from synonymy under pallidefulva (and its synonym F. nitidiventris Emery) by Trager et al. (2007). All Ohio records of this species in Coovert (2005) appear under nitidiventris. This species occurs in mesic, and dry-mesic grasslands, pastures, lawns, old fields, and other open habitats throughout the state. It is undoubtedly more widespread in Ohio than the current records indicate.

integra Nylander. Widespread with majority of the records from unglaciated southern Ohio.

lasioides Emery. Fulton and Lucas Cos. in northwestern Ohio. Rare

montana Wheeler, W.M. Few sites in northcentral Ohio, south to Madison Co.

neogagates Viereck. Scattered sites across northeastern Ohio.

obscuriventris Mayr. Few disjunct sites in extreme northern (Lucas Co.) and southern (Adams, Pike, and Scioto Cos.) Ohio in areas with the appropriate combination of dense woodlands and adjacent open prairie remnants and pastures.

pallidefulva Latreille. Statewide. Distribution of this widespread species in Ohio is confounded with F. incerta (see comments under incerta). A number of records under F. nitidiventris in Coovert (2005) also refer to this species.

pergandei Emery. Few scattered sites in southern and northwestern Ohio.

*podzolica Francoeur. – Summit Co. • 30.57 km SW of Richfield; 41.20992N, 81.66797W; alt. 354 m, 26 July 2007; W. & E. Mackay leg.; W. Mackay det.; MEM221118.

postoculata Kennedy & Dennis. Butler and Hocking Cos. Rare

prociliata Kennedy & Dennis. In Ohio, this species is only known from Ottawa Co. (including type locality) in the extreme northern part of the state. Rare. It is a known inhabitant of glaciated tallgrass prairies and prairie remnants of the upper Midwest.

querquetulana Kennedy & Dennis. Few disjunct sites in extreme southern and northern Ohio where the appropriate combination of open oak woodlands, oak savannas, and associated prairie clearings occurs.

rubicunda Emery. Glaciated western Ohio, and just into the adjacent unglaciated area.

*subaenescens Emery. Literature records: Lucas (Uno et al. 2010). The sole Ohio record appears under the name Formica fusca Linnaeus. As currently understood, true fusca is restricted to the Palearctic region and does not occur in North America (see Schär et al. 2018).

subintegra Wheeler, W.M. Statewide. This dulotic species is found throughout Ohio along with Formica fusca-group hosts.

subsericea Say. Statewide.

ulkei Emery. Widespread in northern Ohio.

vinculans Wheeler, W.M. Scattered sites in glaciated western Ohio, and just into the adjacent unglaciated area.

Polyergus Latreille

lucidus Mayr. This dulotic species is known from few scattered Ohio sites along with host F. incerta.

Tribe Lasiini

Lasius Fabricius

americanus Emery. Statewide. Records under L. alienus (Foerster) in Coovert (2005) refer to this species.

aphidicola (Walsh). Statewide. Records under L. umbratus (Nylander) in Coovert (2005) refer to this species.

brevicornis Emery. Fairfield (OSUC53785, OSUC53786, OSUS53787, OSUC53788). Literature records: Lucas, Montgomery (Uno et al. 2010; Campbell and Crist 2017). This species was previously only known from Summit Co. and an unspecified southcentral Ohio record in Wesson and Wesson (1940). Rare. Records under L. flavus (Fabricius) in Coovert (2005) refer to this species.

claviger (Roger). Widespread with majority of the records from glaciated western Ohio. Under Acanthomyops in Coovert (2005).

interjectus Mayr. Known from a number of scattered Ohio sites, predominantly in the western, glaciated, part of the state. Under Acanthomyops in Coovert (2005).

latipes (Walsh). – Greene Co. • Wright State University Woods; 39.7836N, 84.0636W; 28 July 1998; G.A. Coovert leg.; G.A.C. 2026 #9; G.A. Coovert det.; OSUC57890. Until now, this species was known only from Lucas Co. in glaciated northwestern Ohio (Wing 1968; material at MCZ: MCZENT557430, MCZENT557431, MCZENT557432). Rare. Interestingly, the single specimen in the OSU collection was collected and identified by Gary Coovert but not included in the 2005 list. Under Acanthomyops in Coovert (2005).

minutus Emery. Lucas Co. Rare.

nearcticus Wheeler, W.M. Statewide.

neoniger Emery. Statewide.

speculiventris Emery. Widespread.

Nylanderia Emery

*arenivaga (Wheeler, W.M.). Lucas: Toledo (S. Philpott unpublished data; pers. comm., June 2007). This distinctive yellow Nylanderia is a sand specialist associated with deep sand deposits in dry open areas with sparse vegetation.

faisonensis (Forel). Widespread. Under Paratrechina in Coovert (2005).

*, Eflavipes (Smith). Literature records: Cuyahoga, Franklin, Lake, Lucas (Ivanov and Milligan 2008; Ivanov et al. 2011; Ivanov 2016; Uno et al. 2010). This temperate introduced species is abundant in the urban and suburban areas of Cleveland, Columbus, and Toledo including forested green spaces, gardens, yards, and vacant lots. It is also occasionally found indoors (see Ivanov 2016).

parvula (Mayr). Widespread with majority of the records from southern Ohio. Under Paratrechina in Coovert (2005).

*terricola (Buckley). – Cuyahoga Co. • Cleveland, Cleveland State University campus; 41.50236N, 81.67353W; alt. 202 m; 10 May 2007; K. Ivanov leg.; KI1731; single male on bare soil surface; landscaped flower bed surrounded by pavement; urban; VMNH110553. Workers of this species are nearly impossible to separate from those of N. vividula (Nylander) and males are required for proper identification (see Kallal and LaPolla 2012).

*vividula (Nylander). Cuyahoga (VMNH110554). Literature records: Franklin (Kallal and LaPolla 2012; material at MCZ: MCZENT565308, MCZENT565309, MCZENT565310).

Paratrechina Donisthorpe

*, Elongicornis (Latreille). Literature records: Hamilton (Hedges 1998). In Ohio, this notorious tramp species is currently confined to heated buildings. It is undoubtedly more widespread in the state than current records indicate.

Prenolepis Mayr

imparis (Say). Statewide.

Tribe Myrmelachistini

Brachymyrmex Mayr

depilis Emery. Statewide but uncommonly collected likely due to small worker and colony size, and its largely subterranean habits. This species has an enormous geographic range and may turn out to represent a complex of sibling species.

Subfamily Myrmicinae [15 genera; 65 species, 3 morphospecies]

Tribe Attini

Pheidole Westwood

bicarinata Mayr. Few disjunct sites in extreme northern (Lucas Co.; Friedrich 2010) and southern (Adams, Pike, and Jackson Cos.) Ohio.

Ebilimeki Mayr. The single Ohio record of this introduced species is from a greenhouse in Franklin Co. (Coovert 2005).

*dentata Mayr. – Franklin Co. • Columbus; 7 Apr. 1916; M.R. Smith leg. Single uncatalogued specimen at MCZ (D. Lubertazzi pers. comm., March 2019).

pilifera (Roger). Widespread in southern Ohio.

tysoni Forel. Hocking, Scioto (VMNH110560, VMNH110561, VMNH110596). Literature records: Butler, Montgomery, Preble (Campbell and Crist 2017). Until now, this species was known from Hocking and Gallia Cos. in the southern, unglaciated, part of Ohio.

Strumigenys Smith F.

The perceived rarity of these cryptic litter and soil inhabitants can be explained, at least partly, by the fact that litter extraction techniques historically have been rarely used for the collection of soil and litter dwelling taxa in Ohio. At least some members of this genus (i.e., S. ohioensis Kennedy & Schramm, S. ornata Mayr, S. rostrata Emery) are among the most widespread and commonly encountered ants in the soil and litter layers of eastern deciduous forests. All taxa under Smithistruma in Coovert (2005).

abdita Wesson, L.G. & Wesson, R.G. Literature records: Cuyahoga (Ivanov and Keiper 2010). Until now, this species was known only from Jackson and Ottawa Cos. Rare.

bimarginata Wesson, L.G. & Wesson, R.G. Adams Co. Rare.

clypeata Roger. Literature records: Butler, Montgomery, Preble (Campbell and Crist 2017). Previously only known from Jackson Co. in southcentral Ohio.

dietrichi Smith, M.R. Few scattered sites throughout Ohio.

*hyalina (Bolton). Cuyahoga, Greene (VMNH110584, VMNH110585, VMNH110586, VMNH110587). Literature records: Ottawa (Bolton 2000). Until now, this rarely collected species was only known in Ohio from the type locality in Catawba Beach (Bolton 2000).

*laevinasis Smith, M.R. Greene (VMNH110559). Literature records: Pike (Campbell et al. 2013). This uncommonly collected woodland species is at the northern fringes of its distribution in Ohio.

missouriensis Smith, M.R. Pike Co. Rare.

ohioensis Kennedy & Schramm. Widespread in southern Ohio.

ornata Mayr. Pike Co. Rare.

pergandei Emery. Few scattered sites throughout Ohio.

pilinasis Forel. Pike Co. Rare.

pulchella Emery. Few scattered sites throughout Ohio.

reflexa Wesson, L.G. & Wesson, R.G. Few scattered sites throughout Ohio.

rostrata Emery. Few records from unglaciated southcentral Ohio.

talpa Weber. Few records from unglaciated southcentral Ohio.

Trachymyrmex Forel

septentrionalis (McCook). In Ohio, this wide-ranging fungus-growing ant is known only from Adams and Jackson Cos. in the extreme southern part of the state along the northern fringes of its distribution. Rare.

Tribe Crematogastrini

Cardiocondyla Emery

*, Eobscurior Wheeler, W.M. Literature records: Hamilton (Ivanov 2016). The single Ohio record of this introduced species is from a conservatory in Cincinnati.

Crematogaster Lund

cerasi (Fitch). Statewide.

lineolata (Say). Statewide.

pilosa Emery. Widespread in southcentral Ohio, north to Fairfield Co.

Harpagoxenus Forel

*canadensis Smith, M.R. – Jackson Co. • 1938; L.G. Wesson leg.; FMNHINS113834. In Ohio, this boreal dulotic species is near the southern edge of its known distribution.

Leptothorax Mayr

canadensis Provancher. Ashtabula, Franklin, Geauga (UCMC94034, UCMC94036, VMNH110562, VMNH11563, VMNH110564). Previously only known from Ashtabula Co. in northernmost Ohio. Rare. Records under L. muscorum (Nylander) in Coovert (2005) refer to this species.

Myrmecina Curtis

americana Emery. Statewide.

Temnothorax Mayr

All Temnothorax records, except T. americanus (see below), appear under Leptothorax in Coovert (2005).

ambiguus (Emery). Statewide.

americanus (Emery). This dulotic species is known from a few scattered Ohio sites along with hosts T. ambiguus, T. curvispinosus (Mayr), and T. longispinosus (Roger). Under Protomognathus in Coovert (2005).

curvispinosus (Mayr). Statewide.

duloticus (Wesson, L.G.). This rarely collected dulotic species is known from a handful of scattered Ohio sites along with hosts T. ambiguus, T. curvispinosus, and T. longispinosus.

longispinosus (Roger). Statewide.

minutissimus (Smith, M.R.). In Ohio, this rarely collected workerless parasite of T. curvispinosus is known only from Delaware and Franklin Cos. Rare.

pergandei (Emery). Widespread in southern Ohio.

schaumii (Roger). Statewide.

smithi (Baroni Urbani). This species is only known from Hocking and Jackson Cos. in unglaciated southern Ohio along the northern edge of its distribution. Rare.

texanus (Wheeler, W.M.). Several disjunct records from Jackson (south) and Lucas (north) Cos. Rare.

Tetramorium Mayr

Eatratulum (Schenck). Cuyahoga (CMNHENT40727, CMNHENT40728), Preble (K. Campbell unpublished data; pers. comm., February 2019). Literature records: Ashtabula, Cuyahoga, Geauga (Ivanov 2016). This introduced workerless social parasite of Tetramorium immigrans Santschi previously was only known from Shelby Co. in western Ohio. It is rarely observed in both its native and introduced ranges, presumably because of its parasitic lifestyle or simply because it is indeed rare. Under Anergates in Coovert (2005).

*,Ebicarinatum (Nylander). Literature records: Franklin: Columbus, 1931, collector unknown, material at the US National Museum of Natural History (Wetterer 2009), Ohio (Bolton 1979). In temperate areas, this widely distributed tramp species is confined to greenhouses and other heated buildings.

Eimmigrans Santschi. Statewide. This introduced species is abundant in disturbed habitats and urban settings throughout Ohio, and is also occasionally found indoors. Records under T. caespitum (Linnaeus) in Coovert (2005) refer to this species.

Tribe Myrmicini

Myrmica Latreille

André Francoeur is working on a much-needed revision of the Nearctic members of this genus and I have chosen to follow his preliminary findings in my species accounts below including the recognition of a few as-yet-undescribed species (see also Ellison et al. 2012).

americana Weber. Distribution of this northerly species in Ohio is unclear, confounded with the undescribed and more widely distributed M. AF-eva. True americana appears to be a strict psammophile occurring in glaciated sand communities and probably does not occur outside of them (J. Trager pers. comm.). In Ohio, I have only collected this species in the sparsely vegetated fossil dune ridge of North Kingsville Sand Barrens in Ashtabula Co. in northernmost Ohio (VMNH110565). It likely also occurs in other sand based communities along the coast of Lake Erie in northern Ohio, however, its distribution in the state is much more restricted than indicated in Coovert (2005).

emeryana Forel. Distribution of this species in Ohio is unclear, confounded with the undescribed but widespread and abundant M. AF-scu and M. AF-smi. The distribution depicted in Coovert (2005) represents a composite of the distributions of these three taxa. Records under Myrmica latifrons Stärcke in Coovert (2005) refer to this species.

*AF-eva. – Cuyahoga Co. • Cleveland Heights, 3986 Bluestone Rd.; 41.52998N, 82.53670W; 30 Sep. 2004; H. Clebsch leg.; yellow pan traps; CMNHENT0038652. – Hardin Co. • Ada, Ohio Northern University campus; 40.76844N, 83.84341W; alt. 294 m; 8–9 Sep. 2017; A. Brooks & J. Isaak leg.; red pan traps; mowed grass at edge of a small woodlot; VMNH110577 • same collection data as for preceding; VMNH110578. – Hocking Co. • Hocking Hills, Deep Woods farm; 39.40812N, 82.57503W; alt. 215 m; 14 June 2008; K. Ivanov leg.; KI2042; ground foragers; open mowed grass field; VMNH110545. – Madison Co. • OPOTA tactical training center; 39.88028N, 83.48917W; alt. 338 m; 25–26 July 2016; J.B. Keiper leg.; old field; sweeping; VMNH110579 • same collection data as for preceding; VMNH110580. – Summit Co. • Sagamore Hills, Eaton estate; 20 Sep. 2002; M.R. Proffitt leg.; CMNHENT0038653 • Singer Lake Bog; 40.92288N, 81.48720W; alt. 356 m; 9 May 2008; K. Ivanov leg.; KI2019; ground foragers; edge of agricultural field; VMNH110546. This undescribed species is found in dry-mesic to mesic grasslands, pastures, and old fields (Ellison et al. 2012). Historically it has been commonly confused with Myrmica americana.

fracticornis Forel. Statewide, in moist open habitats. Historical records likely confounded with the undescribed M. AF-scu.

*incompleta Provancher. Franklin (OSUC270669; as M. brevinodis Emery, N.F. Johnson det. 1984). Literature records: Adams (Weber 1950). This species was prematurely excluded from the 2005 list without examination of Weber’s specimens.

*lobifrons Pergande. – Champaign Co. • Cedar Bog Nature Preserve; 40.05785N, 83.79613W; 27– 28 May 2006; T. Jones leg.; sedge meadow; VMNH110548. – Cuyahoga Co. • Cleveland Metroparks, Acacia Reservation; 41.51039N, 81.49517W; alt. 313 m; 27 May 2013; K. Ivanov leg.; KI2333; sedge meadow; sweeping; VMNH110547. – Stark Co. • Jackson Township, Willow Dale Bog; 22 Sep. 2011; D. Kriska leg.; VMNH110549 • same collection data as for preceding; VMNH110550 • same collection data as for preceding; VMNH110551 • same collection data as for preceding; VMNH110552. In Ohio, this boreal bog-inhabiting species is rarely collected in the northern, glaciated, part of the state.

pinetorum Wheeler, W.M. Statewide.

punctiventris Roger. Statewide.

*AF-scu. – Ashtabula Co. • North Kingsville Sand Barrens; 30 Aug. – 24 Sep. 2007; T. Pucci leg.; Malaise trap; CMNHENT0038834 • same collection data as for preceding; 9– 19 Sep. 2008; VMNH110603 • Pymatuning Creek Fen; 41.56738N, 80.62710W; alt. 298 m; 13 Oct. 2008; K. Ivanov leg.; KI2192; soil nest under plant litter at the base of a small sumac (Rhus sp.); edge of agricultural field; VMNH110598. – Cuyahoga Co. • Strongsville, Cleveland Metroparks, nr. Greenbrier L.; 8 Aug. 2009; T. Pucci leg.; meadow; sweeping; CMNHENT0038835. – Erie Co. • Kelleys Island, Coleman Tract; 2–4 Aug. 2002; B. Coleman leg.; C1; pitfall trap; VMNH110571 • same collection data as for preceding; D2; VMNH110569 • same collection data as for preceding; 6–8 Sep. 2002; D2; VMNH110570 • Castalia, Rockwell Springs Trout Club; 41.39014N, 82.83646W; alt. 194 m; 17 July 2008; K. Ivanov leg.; KI2121; strays in grass litter; open grass field adjacent to riparian woods; VMNH110600 • Castalia, 0.25 mi N intersection Rt. 312 and Vickery Rd.; 41.38979N, 82.84770W; alt. 192 m; 17 July 2008; K. Ivanov leg.; KI2124; ground foragers; managed tallgrass prairie remnant; VMNH110599 – Geauga Co. • Burton Wetlands Nature Preserve; 41.44576N, 81.17723W; alt. 341 m; 21 Apr. 2006; K. Ivanov leg.; KI1228; foragers in grass litter at base of a large oak tree; edge of an open grass field; VMNH110590 • Fern Lake Bog; 41.44026N, 81.17262W; alt. 344 m; 16 May 2007; K. Ivanov leg.; KI1745; soil nest in clay loam under grass litter; edge of tamarack bog; VMNH110589 • Taber Preserve; 18 May 2007; T. Pucci leg.; sweeping; CMNHENT0038833 • Soubusta; 41.57000N, 81.24000W; 28 July 2009; T.H. Webster leg.; scrap wood pile; CMNHENT0038816 – Greene Co. • Wright State University campus; 8 July 2009; B. Doane leg.; tuna baits; VMNH110605. – Madison Co. • OPOTA tactical training center; 39.88028N, 83.48917W; alt. 338 m; 25–26 July 2016; J.B. Keiper leg.; old field; sweeping; VMNH110581 • same collection data as for preceding VMNH110582. – Medina Co. • Wolf Creek Environmental Center; 2 July 2008; T. Pucci leg.; sweeping; CMNHENT0038815. – Ottawa Co. • Winous Point; 24 June 2004; T. Pucci leg.; sweeping; CMNHENT0038814 • Ottawa National Wildlife Refuge, Young Pond; 41.53649N, 83.00608W; alt. 175 m; 25 July 2008; K. Ivanov leg.; KI2147; ground foragers; open riparian woods; VMNH110568. – Trumbull Co. • Chamberlin Forest; 41.44773N, 80.98881W; alt. 311 m; 5 June 2009; K. Ivanov leg.; KI2201; ground foragers; small grass field surrounded by mesic woods; VMNH110567. – Wyandot Co. • Kildeer Plains Wildlife Area; 40.70234N, 83.24097W; alt. 271 m; 22 May 2009; K. Ivanov leg.; KI2197; ground foragers; open grass and gravel road at the edge of an open mixed mesic woodland; VMNH110591. This undescribed species is found in mesic grasslands, wood’s edges, and open, remnant dry-mesic and mesic woodlands. Historically it has been confused with M. emeryana and M. fracticornis. It is undoubtedly more widespread in the state than the above records indicate.

*semiparasitica Francoeur. Literature records: Cuyahoga (Francoeur and Ivanov 2008; Ivanov and Keiper 2009, 2011; Ivanov et al. 2010). This recently described species has been rarely collected in the northern half of the eastern deciduous forest biome along with its suspected host M. punctiventris. Since the range of its host covers the southern half of the eastern deciduous forest biome as well, it is expected that M. semiparasitica will be discovered much further south in the future

*AF-smi. Athens, Cuyahoga, Erie, Geauga, Greene, Hocking, Lake, Medina, Scioto, Summit (CMNHENT0038654, CMNHENT0038655, CMNHENT0038656, CMNHENT0038657, CMNHENT0038658, CMNHENT0038659, CMNHENT0038660, CMNHENT0038661, CMNHENT0038662, CMNHENT0038663, CMNHENT0038664, CMNHENT0038665, CMNHENT0038666, CMNHENT0038667, CMNHENT0038668, CMNHENT0038669, CMNHENT0038670, CMNHENT0038671, CMNHENT0038672, CMNHENT0038673, CMNHENT0038674, CMNHENT0038675, CMNHENT0038676, CMNHENT0038677, CMNHENT0038678, CMNHENT0038679, CMNHENT0038680, CMNHENT0038681, CMNHENT0038682, CMNHENT0038683, CMNHENT0038684, CMNHENT0038685, CMNHENT0038687, CMNHENT0038688, CMNHENT0038811, CMNHENT0038812, CMNHENT0038813, CMNHENT0038817, CMNHENT0038819, CMNHENT0038820, CMNHENT0038838, CMNHENT0038839, CMNHENT0038840, VMNH110572, VMNH110573, VMNH110574, VMNH110583, VMNH110588, VMNH110592, VMNH110594, VMNH110595, VMNH110604). Literature records: Cuyahoga, Medina (Ivanov and Keiper 2009, 2010, 2011; Ivanov et al. 2010). This undescribed species prefers dry to mesic deciduous woodlands and mixed forests (Ellison et al. 2012) and is also occasionally found in mesic grasslands and old fields. It is undoubtedly more widespread in the state than the above records indicate.

Tribe Solenopsidini

Monomorium Mayr

minimum (Buckley). Statewide but less common in northern Ohio.

Epharaonis (Linnaeus). This notorious tramp species is known from a number of scattered indoor records across Ohio.

Solenopsis Westwood

Members of the molesta complex (sensu Pacheco and Mackay 2013) are minute in size and notoriously difficult to identify. Historical Ohio records of this complex (i.e., S. carolinensis Forel, S. molesta (Say), and S. texana Emery) are likely confounded with each other and should be viewed with caution. Presence of queens will aid in identification, but see discussion in Deyrup (2016).

carolinensis Forel. Single unspecified southcentral Ohio record in Wesson and Wesson (1940). Rare.

molesta (Say). Statewide.

texana Emery. Adams Co. Rare.

Tribe Stenammini

Aphaenogaster Mayr

While many eastern Aphaenogaster can be readily identified based on morphological characters, a number of taxa remain difficult, or impossible, to separate based on morphology alone. This is especially true for some members of the rudis clade although some progress has been recently made (DeMarco 2015; DeMarco and Cognato 2016). Historical records of Aphaenogaster carolinensis Wheeler, W.M. and Aphaenogaster rudis Enzmann from Ohio, and elsewhere, should be interpreted with caution.

fulva Roger. Statewide.

lamellidens Mayr. An unspecified southern Ohio record in Dennis (1938). Rare.

mariae Forel. A number of scattered sites in unglaciated southern Ohio.

picea (Wheeler, W.M.). Statewide.

rudis Enzmann. Statewide.

tennesseensis (Mayr). Statewide.

treatae Forel. Few scattered records from prairie remnants and old fields in extreme northern and southern Ohio.

Stenamma Westwood

brevicorne (Mayr). Statewide.

*diecki Emery. – Licking Co. • Blackhand Gorge; 3–4 May 1989; P. Kovarik leg.; berlesate; leaf litter; S.P. Cover det. 1989; MCZ540382 • same collection data as for preceding; MCZ540383.

impar Forel. Widespread and likely more common than current records indicate.

schmitti Wheeler, W.M. Few scattered sites throughout Ohio.

Subfamily Ponerinae [3 genera; 4 species]

Tribe Ponerini

Brachyponera Emery

*, Echinensis (Emery). Literature records: Hamilton: Cincinnati, 2017, J Boggs leg., Ohio State University Extension (Guénard et al. 2018). This East Asian species has been spreading through the eastern United States since its original discovery in Georgia, North Carolina, and Virginia (Smith 1934). At present, Ohio lies along the northern edge of its known introduced range. Given its profound negative impacts on local insect populations, care should be taken in monitoring the future spread of this invasive species in Ohio and the adjacent areas.

Hypoponera Santschi

opacior (Forel). Widespread in southern Ohio, north to Guernsey Co.

Eragusai (Emery). This introduced species in known from a single locality in Jackson Co. (Wesson and Wesson 1940). Record under H. gleadowi (Emery) in Coovert (2005) refers to this species.

Ponera Latreille

pennsylvanica Buckley. Statewide.

Subfamily Proceratiinae [1 genus; 3 species]

Tribe Proceratiini

Proceratium Roger

*crassicorne Emery. Literature records: southcentral Ohio (Wesson and Wesson 1940). Rare. The single Ohio record is included under P. silaceum in Coovert (2005). This species was revived from synonymy under P. silaceum by Baroni Urbani and De Andrade (2003).

pergandei (Emery). Literature records: Franklin (Campbell et al. 2013). Until now, this species was only known from Adams and Greene Cos. Rare.

silaceum Roger. Few scattered sites throughout Ohio.

Doubtful Ohio records

Dorymyrmex insanus (Buckley) (Friedrich 2010; likely misidentification of D. grandulus), Pheidole tepicana Pergande (TTU-Z_075141), Pogonomyrmex barbatus (Smith F.) (OSUC50490), and Leptogenys elongata (Buckley) (INHS-INSECT-575849) are herein excluded from the Ohio list. These records represent stark distribution anomalies relative to the rest of their known western ranges and are likely based on misidentifications and/or mislabeled specimens. The record of A. carolinensis from Ottawa Co. reported in Mackay and Mackay (2017; p.238) seems rather surprising. The finding of this southern species in northern Ohio is doubtful and likely represents misidentification of the more common and widespread A. rudis. Workers from young A. rudis colonies can easily be mistaken for A. carolinensis and genetic data is needed for proper identification (see DeMarco and Cognato 2016). I provisionally exclude this species from the Ohio list awaiting future work on the rudis Complex of species. In addition, Mackay and Mackay (2017) erroneously cite Coovert (2005) as source of Aphaenogaster ashmeadi (Emery) (p. 211) and A. flemingi Smith, M.R. (p. 244) records from Ohio. To my knowledge, no Ohio records of these two southeastern taxa exist.

Acknowledgements

I thank J. Bissell, K. Campbell, H. Clebsch, B. Coleman, B. Doane, R. Friedrich, M. Gates, R. Gibson, S. Heideman, J. Keiper, D. Kriska, W. Mackay, A. Perez, S. Philpott, B. Patrick, B. Poynter, T. Pucci, M. Vincent, and T. Webster for providing specimens and/or information concerning the distribution of Ohio ants. I also thank G. Coovert for sharing his knowledge of Ohio ants during the early stages of this work. I thank J. MacGown (MEM) for verifying Ohio records at the Mississippi Entomological Museum, D. Lubertazzi (MCZ) for verifying P. dentata record at the Louis Agassiz Museum of Comparative Zoology, and G. Svenson and N. Gunter (CMNH) for providing accommodations and access to the invertebrate collection at CMNH, and for the loan of specimens. Comments and suggestions from O. Lockhart, J. Trager, and two anonymous reviewers improved the manuscript. This research was partially supported by the Cleveland State University DDRE award program, and the Ohio Biological Survey and Geauga County Park District grant programs.

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Appendix 1

Alphabetical list of native ant taxa documented from surrounding areas but not yet recorded from Ohio. Data from Wheeler et al. (1994), Coovert (2005), Carroll (2011), Seifert et al. (2014), Butler and Coulter (2019), Antmaps (2019), and references therein.

Species Indiana Kentucky Michigan Pennsylvania West Virginia
Camponotus decipiens Emery
Crematogaster ashmeadi Mayr
Crematogaster laeviuscula Mayr
Crematogaster missouriensis Emery
Dolichoderus mariae Forel
Formica adamsi Wheeler, W.M.
Formica creightoni Buren
Formica ferocula Wheeler, W.M
Formica fossaceps Buren
Formica hewitti Wheeler, W.M.
Formica impexa Wheeler, W.M.
Formica indianensis Cole
Formica neorufibarbis Emery
Formica nepticula Wheeler, W.M.
Formica obscuripes Forel
Formica talbotae Wilson
Formicoxenus hirticornis (Emery)
Formicoxenus provancheri (Emery)
Lasius murphyi Forel
Lasius pallitarsis (Provancher)
Lasius plumopilosus Buren
Lasius subglaber Emery
Lasius subumbratus Viereck
Monomorium emarginatum DuBois
Monomorium talbotae DuBois
Myrmica alaskensis Wheeler, W.M.
Myrmica brevispinosa Wheeler, W.M.
Myrmica detritinodis Emery
Myrmica monticola Creighton
Myrmica nearctica Weber
Myrmica spatulata Smith, M.R.
Neivamyrmex nigrescens (Cresson)
Pheidole morrisii Forel
Polyergus bicolor Wasmann
Polyergus breviceps Emery
Polyergus mexicanus Forel
Polyergus montivagus Wheeler, W.M.
Polyergus sanwaldi Trager
Stenamma meridionale Smith, M.R.
Strumigenys angulata Smith, M.R.
Strumigenys filitalpa (Brown)
Strumigenys memorialis (Deyrup)
Strumigenys metazytes (Bolton)
Temnothorax pilagens Seifert et al.