United States

Department of 

Agriculture

Forest Service

Northern 

Research Station

General Technical

Report NRS-16

Spiders (Arachnida: Araneae) of Milbridge, 

Washington County, Maine

Daniel T. Jennings

Frank Graham, Jr.
Visit our homepage at: http://www.nrs.fs.fed.us/

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USDA FOREST SERVICE USDA Forest Service 

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 Delaware, OH 43015-8640

October 2007 Fax: (740)368-0152

The use of trade, firm, or corporation names in this publication is for the information and 

convenience of the reader. Such use does not constitute an official endorsement or approval 

by the U.S. Department of Agriculture, Forest Service, of any product or service to the 

exclusion of others that may be suitable.

ABSTRACT

Adult spiders (N = 6,979) of 19 families, 145 genera, and 302 species (including 4 unknown) 

were collected from diverse inland, coastal, and offshore habitats of Milbridge, a 6,290-ha 

minor civil division of the East Coastal BioPhysical Region. Spider species richness per 

family ranged from 1 (Titanoecidae and Pisauridae) to 89 (Linyphiidae); species richness per 

genus ranged from 1 to 13, with 88 genera represented by a single species. As expected, the 

collected taxa were distributed unequally between two basic foraging strategies: 10 families, 

98 genera, and 179 species of web spinners; 9 families, 47 genera, and 123 species of 

hunters.

Spider abundances varied widely among taxa; individuals per family ranged from 1 

(Titanoecidae and Pisauridae) to 1,691(Lycosidae); individuals per species ranged from 1 

(86 species) to 470. Pardosa moesta was the most frequently collected spider. Although total 

species composition favored web spinners over hunters, more hunters were collected than 

web spinners, and more female spiders were collected than male spiders, a pattern evident 

for both web spinners and hunters. Spider sex ratios varied widely among the collected 

species and were influenced by sampling method, habitat, and season. Pitfall traps yielded 

more species and more individuals than any other sampling method. Fully 47.0 percent of the 

inventoried fauna were method-unique species; most were taken by pitfall traps, searches, 

and sweep nets. Species-faunal compositions among habitats were generally distinct among 

habitats (QS ≤ 50.0), and included habitat-unique spedies. 

Species-range extensions include 31 new state records, 6 New England records, and 3 

national records. At least 12 invasive species were among the inventoried spiders; 5 were 

associated with domestic habitats. Based on published and unpublished records from the 

same biophysical region, total estimated richness is 24 families, 172 genera, and ~ 411 

species. Loss of habitat and displacement of native species by invasives pose potential 

threats to this diverse fauna.

COVER PHOTO

Araneus orb weaver, courtesy of Lt. Col. Murray Cragin, USAF (retired).
CONTENTS

Introduction......................................................................................................................1

Study Sites and Habitats ................................................................................................2

 Study Sites ....................................................................................................................2

 Habitats .........................................................................................................................2

Materials and Methods ....................................................................................................2

 Spider Collection Methods .............................................................................................2

 Pitfall traps ................................................................................................................2

 Sweep nets ...............................................................................................................5

 Beating sheets ..........................................................................................................5

 Tree-bole brush and net ...........................................................................................5

 Litter condenser-extraction .......................................................................................5

 Searches ..................................................................................................................5

 Collection Site and Habitat Descriptors .........................................................................6

 Global positioning system coordinates .....................................................................6

 Habitat descriptors ....................................................................................................6

 Specimen Identifi cations and Museum Deposits ...........................................................6

 Spider identifi cations ................................................................................................6

 Museum specimen deposits .....................................................................................6

 Data Summaries and Analyses .....................................................................................6

 Data summaries .......................................................................................................6

 Data analyses ...........................................................................................................7

Annotated Faunal List: Introduction............................................................................ 11

Annotated Faunal List: Web Spinner Families ...........................................................12

 Family Pholcidae .........................................................................................................12

 Pholcus manueli Gertsch, 1937 ..............................................................................12

 Family Theridiidae .......................................................................................................13

 Achaearanea globosa (Hentz, 1850) ......................................................................13

 Achaearanea tabulata Levi, 1980 ...........................................................................13

 Crustulina sticta (O. Pickard-Cambridge, 1861) .....................................................14

 Dipoena nigra (Emerton, 1882) ..............................................................................15

 Enoplognatha intrepida (Sørensen, 1898) ..............................................................15

 Enoplognatha ovata (Clerck, 1757) ........................................................................15

 Euryopis funebris (Hentz, 1850) .............................................................................16

 Neospintharus trigonum (Hentz, 1850) ...................................................................17

 Neottiura bimaculata (Linnaeus, 1767) ...................................................................17

 Phoroncidia americana (Emerton, 1882) ................................................................18

 Robertus banksi (Kaston, 1946) .............................................................................18

 Rugathodes aurantius (Emerton, 1915) .................................................................18

 Rugathodes sexpunctatus (Emerton, 1882) ...........................................................19

 Steatoda americana (Emerton, 1882) ....................................................................19
 Steatoda bipunctata (Linnaeus, 1758) ....................................................................19

 Steatoda borealis (Hentz, 1850) .............................................................................20

 Takayus lyricus (Walckenaer, 1842) .......................................................................20

 Theridion albidum Banks, 1895 ..............................................................................21

 Theridion differens Emerton, 1882 .........................................................................21

 Theridion frondeum Hentz, 1850 ............................................................................21

 Theridion glaucescens Becker, 1879 ......................................................................22

 Theridion montanum Emerton, 1882 ......................................................................22

 Theridion murarium Emerton, 1882 ........................................................................23

 Theridion pictum (Walckenaer, 1802) .....................................................................23

 Theridula emertoni Levi, 1954 ................................................................................24

 Thymoites unimaculatus (Emerton, 1882) ..............................................................24

 Wamba crispulus (Simon, 1895) .............................................................................25

 Family Linyphiidae—Subfamily Linyphiinae ................................................................25

 Allomengea dentisetis (Grube, 1861) .....................................................................26

 Bathyphantes brevis (Emerton, 1911) ....................................................................26

 Bathyphantes gracilis (Blackwall, 1841) .................................................................26

 Bathyphantes pallidus (Banks, 1892) .....................................................................26

 Centromerus cornupalpis (O. Pickard-Cambridge, 1875) ......................................27

 Centromerus longibulbus (Emerton, 1882) .............................................................27

 Centromerus persolutus (O. Pickard-Cambridge, 1875) ........................................28

 Centromerus sylvaticus (Blackwall, 1841) ..............................................................28

 Diplostyla concolor (Wider, 1834) ...........................................................................28

 Drapetisca alteranda Chamberlin, 1909 .................................................................29

 Estrandia grandaeva (Keyserling, 1886) ................................................................29

 Frontinella communis (Hentz, 1850) .......................................................................29

 Incestophantes calcaratus (Emerton, 1909) ...........................................................30

 Incestophantes duplicatus (Emerton, 1913) ...........................................................31

 Kaestneria pullata (O. Pickard-Cambridge, 1863) ..................................................31

 Lepthyphantes alpinus (Emerton, 1882) .................................................................32

 Lepthyphantes leprosus (Ohlert, 1865) ..................................................................32

 Lepthyphantes turbatrix (O. Pickard-Cambridge, 1877) .........................................32

 Linyphia triangularis (Clerck, 1757) ........................................................................33

 Macrargus multesimus (O. Pickard-Cambridge, 1875) ..........................................33

 Meioneta fabra (Keyserling, 1886) .........................................................................34

 Meioneta nigripes (Simon, 1884) ............................................................................34

 Meioneta simplex (Emerton, 1926) .........................................................................35

 Meioneta unimaculata (Banks, 1892) .....................................................................35

 Meioneta zygia (Keyserling, 1886) .........................................................................36

 Microlinyphia mandibulata (Emerton, 1882) ...........................................................36
 Microneta viaria (Blackwall, 1841) ..........................................................................36

 Neriene clathrata (Sundevall, 1830) .......................................................................37

 Neriene radiata (Walckenaer, 1842) .......................................................................37

 Pityohyphantes costatus (Hentz, 1850) ..................................................................38

 Pityohyphantes phrygianus (C. L. Koch, 1836) ......................................................38

 Porrhomma convexum (Westring, 1851) ................................................................39

 Porrhomma terrestre (Emerton, 1882) ...................................................................39

 Tapinopa bilineata Banks, 1893 ..............................................................................40

 Tenuiphantes zebra (Emerton, 1882) .....................................................................40

 Family Linyphiidae—Subfamily Erigoninae .................................................................41

 Baryphyma trifrons (O. Pickard-Cambridge, 1863) ................................................41

 Carorita limnaea (Crosby & Bishop, 1927) .............................................................42

 Ceraticelus bulbosus (Emerton, 1882) ...................................................................42

 Ceraticelus emertoni (O. Pickard-Cambridge, 1874) ..............................................42

 Ceraticelus fi ssiceps (O. Pickard-Cambridge, 1874) ..............................................43

 Ceraticelus laetabilis (O. Pickard-Cambridge, 1874) ..............................................43

 Ceraticelus laticeps (Emerton, 1894) .....................................................................43

 Ceraticelus minutus (Emerton, 1882) .....................................................................44

 Ceratinella brunnea Emerton, 1882 ........................................................................44

 Ceratinella parvula (Fox, 1891) ..............................................................................44

 Ceratinops annulipes (Banks, 1892) ......................................................................44

 Ceratinopsis nigriceps Emerton, 1882 ....................................................................45

 Collinsia plumosa (Emerton, 1882) ........................................................................45

 Diplocephalus cristatus (Blackwall, 1833) ..............................................................46

 Diplocephalus subrostratus (O. Pickard-Cambridge, 1873) ...................................46

 Dismodicus decemoculatus (Emerton, 1882) .........................................................46

 Eperigone entomologica (Emerton, 1911) ..............................................................47

 Eperigone undulata (Emerton, 1914) .....................................................................47

 Erigone aletris Crosby & Bishop, 1928 ...................................................................47

 Erigone atra Blackwall, 1833 ..................................................................................47

 Erigone blaesa Crosby & Bishop, 1928 ..................................................................48

 Erigone dentipalpis (Wider, 1834) ..........................................................................48

 Erigone ephala Crosby & Bishop, 1928 ..................................................................49

 Erigone zographica Crosby & Bishop, 1928 ...........................................................49

 Floricomus rostratus (Emerton, 1882) ....................................................................49

 Gonatium crassipalpum Bryant, 1933 ....................................................................49

 Grammonota angusta Dondale, 1959 ....................................................................50

 Grammonota gigas (Banks, 1896) ..........................................................................51

 Grammonota maritima Emerton, 1925 ...................................................................51

 Grammonota trivittata Banks, 1895 ........................................................................51
 Hypselistes fl orens (O. Pickard-Cambridge, 1875) ................................................52

 Islandiana fl aveola (Banks, 1892) ..........................................................................53

 Metopobactrus prominulus (O. Pickard-Cambridge, 1872) ....................................53

 Pocadicnemis americana Millidge, 1976 ................................................................53

 Satilatlas marxi Keyserling, 1886 ...........................................................................54

 Sciastes truncatus (Emerton, 1882) .......................................................................54

 Scotinotylus exsectoides Millidge, 1981 .................................................................54

 Sisicottus montanus (Emerton, 1882) ....................................................................55

 Sisicus penifusifer Bishop & Crosby, 1938 .............................................................55

 Soulgas corticarius (Emerton, 1909) ......................................................................55

 Styloctetor stativus (Simon, 1881) ..........................................................................56

 Tapinocyba simplex (Emerton, 1882) .....................................................................56

 Tapinocyba sp. ........................................................................................................56

 Tunagyna debilis (Banks, 1892) .............................................................................57

 Walckenaeria atrotibialis O. Pickard-Cambridge, 1878 ..........................................57

 Walckenaeria communis (Emerton, 1882) ..............................................................57

 Walckenaeria digitata (Emerton, 1913) ..................................................................57

 Walckenaeria directa (O. Pickard-Cambridge, 1878) .............................................58

 Walckenaeria exigua Millidge, 1983 .......................................................................58

 Walckenaeria lepida (Kulczyński, 1885) .................................................................58

 Walckenaeria redneri Millidge, 1983 .......................................................................59

 Walckenaeria spiralis (Emerton, 1882) ...................................................................59

 Walckenaeria subspiralis Millidge, 1983 .................................................................59

 Undetermined genus, species ................................................................................60

 Family Tetragnathidae .................................................................................................61

 Meta ovalis (Gertsch, 1933) ...................................................................................61

 Pachygnatha autumnalis Marx, 1884 .....................................................................61

 Pachygnatha brevis Keyserling, 1884 ....................................................................62

 Tetragnatha caudata Emerton, 1884 ......................................................................62

 Tetragnatha elongata Walckenaer, 1842 ................................................................62

 Tetragnatha extensa (Linnaeus, 1758) ...................................................................63

 Tetragnatha guatemalensis O. Pickard-Cambridge, 1889 ......................................63

 Tetragnatha laboriosa Hentz, 1850 .........................................................................63

 Tetragnatha straminea Emerton, 1884 ...................................................................64

 Tetragnatha versicolor Walckenaer, 1842 ...............................................................64

 Family Araneidae .........................................................................................................66

 Acanthepeira stellata (Walckenaer, 1805) ..............................................................66

 Araneus cavaticus (Keyserling, 1882) ....................................................................66

 Araneus iviei (Archer, 1951) ...................................................................................67

 Araneus juniperi (Emerton, 1884) ...........................................................................67
 Araneus marmoreus Clerck, 1757 ..........................................................................67

 Araneus nordmanni (Thorell, 1870) ........................................................................68

 Araneus pratensis (Emerton, 1884) ........................................................................69

 Araneus saevus (L. Koch, 1872) ............................................................................69

 Araneus trifolium (Hentz, 1847) ..............................................................................69

 Araniella displicata (Hentz, 1847) ...........................................................................70

 Argiope aurantia (Lucas, 1833) ..............................................................................70

 Argiope trifasciata (Forskål, 1775) ..........................................................................71

 Cyclosa conica (Pallas, 1772) ................................................................................71

 Eustala anastera (Walckenaer, 1842) .....................................................................72

 Eustala cepina (Walckenaer, 1842) ........................................................................72

 Hypsosinga funebris (Keyserling, 1892) .................................................................72

 Hypsosinga pygmaea (Sundevall, 1831) ................................................................73

 Hypsosinga rubens (Hentz, 1847) ..........................................................................73

 Larinioides cornutus (Clerck, 1757) ........................................................................73

 Larinioides patagiatus (Clerck, 1757) .....................................................................74

 Mangora placida (Hentz, 1847) ..............................................................................74

 Metepeira palustris Chamberlin & Ivie, 1942 ..........................................................74

 Neoscona arabesca (Walckenaer, 1842) ................................................................75

 Zygiella atrica (C. Koch, 1845) ...............................................................................76

 Family Agelenidae .......................................................................................................77

 Agelenopsis actuosa (Gertsch & Ivie, 1936) ..........................................................77

 Agelenopsis potteri (Blackwall, 1846) .....................................................................78

 Agelenopsis utahana (Chamberlin & Ivie, 1933) ....................................................78

 Tegenaria domestica (Clerck, 1757) .......................................................................79

 Family Hahniidae .........................................................................................................80

 Antistea brunnea (Emerton, 1909) .........................................................................80

 Cryphoeca montana Emerton, 1909 .......................................................................80

 Hahnia cinerea Emerton, 1890 ...............................................................................81

 Neoantistea agilis (Keyserling, 1887) .....................................................................81

 Neoantistea magna (Keyserling 1887) ...................................................................82

 Family Dictynidae ........................................................................................................83

 Argenna obesa Emerton, 1911 ...............................................................................83

 Cicurina arcuata Keyserling, 1887 ..........................................................................84

 Cicurina brevis (Emerton, 1890) .............................................................................84

 Cicurina pallida Keyserling, 1887 ...........................................................................84

 Cicurina placida Banks, 1892 .................................................................................85

 Dictyna brevitarsa Emerton, 1915 ..........................................................................85

 Dictyna coloradensis Chamberlin, 1919 .................................................................85

 Dictyna minuta Emerton, 1888 ...............................................................................86
 Emblyna manitoba (Ivie, 1947) ...............................................................................86

 Emblyna maxima (Banks, 1892) .............................................................................86

 Emblyna phylax (Gertsch & Ivie, 1936) ..................................................................87

 Emblyna sublata (Hentz, 1850) ..............................................................................87

 Lathys pallida (Marx, 1891) ....................................................................................88

 Family Amaurobiidae ...................................................................................................89

 Amaurobius borealis Emerton, 1909 ......................................................................89

 Callobius bennetti (Blackwall, 1846) .......................................................................89

 Coras montanus (Emerton, 1890) ..........................................................................90

 Cybaeopsis euopla (Bishop & Crosby, 1935) .........................................................90

 Wadotes calcaratus (Keyserling, 1887) ..................................................................91

 Family Titanoecidae .....................................................................................................92

 Titanoeca americana Emerton, 1888......................................................................92

Annotated Faunal List: Hunter Families......................................................................93

 Family Lycosidae .........................................................................................................93

 Alopecosa aculeata (Clerck, 1757) .........................................................................93

 Arctosa emertoni Gertsch, 1934 .............................................................................94

 Arctosa rubicunda (Keyserling, 1877) ....................................................................94

 Hogna frondicola (Emerton, 1885) .........................................................................94

 Pardosa distincta (Blackwall, 1846) ........................................................................95

 Pardosa fuscula (Thorell, 1875) .............................................................................96

 Pardosa groenlandica (Thorell, 1872) ....................................................................96

 Pardosa hyperborea (Thorell, 1872) .......................................................................97

 Pardosa lapidicina Emerton, 1885 ..........................................................................97

 Pardosa mackenziana (Keyserling, 1877) ..............................................................97

 Pardosa modica (Blackwall, 1846) .........................................................................98

 Pardosa moesta Banks, 1892 ................................................................................99

 Pardosa saxatilis (Hentz, 1844) ............................................................................100

 Pardosa xerampelina (Keyserling, 1877) .............................................................100

 Pirata cantralli Wallace & Exline, 1978 .................................................................101

 Pirata insularis Emerton, 1885 .............................................................................102

 Pirata minutus Emerton, 1885 ..............................................................................102

 Pirata piraticus (Clerck, 1757) ..............................................................................102

 Pirata sp. (nr. montanus Emerton, 1885) ..............................................................103

 Schizocosa communis (Emerton, 1885) ...............................................................103

 Schizocosa crassipalpata Roewer, 1951 ..............................................................104

 Schizocosa saltatrix (Hentz, 1844) .......................................................................104

 Trabeops aurantiacus (Emerton, 1885) ................................................................104

 Trochosa ruricola (De Geer, 1778) .......................................................................105

 Trochosa terricola Thorell, 1856 ...........................................................................105
 Family Pisauridae ......................................................................................................107

 Dolomedes striatus Giebel, 1869 .........................................................................107

 Family Liocranidae ....................................................................................................108

 Agroeca ornata Banks, 1892 ................................................................................108

 Agroeca pratensis Emerton, 1890 ........................................................................108

 Family Clubionidae ....................................................................................................109

 Clubiona bishopi Edwards, 1958 ..........................................................................109

 Clubiona bryantae Gertsch, 1941 ......................................................................... 110

 Clubiona canadensis Emerton, 1890 .................................................................... 110

 Clubiona johnsoni Gertsch, 1941 ......................................................................... 111

 Clubiona kastoni Gertsch, 1941 ........................................................................... 111

 Clubiona littoralis Banks, 1895 ............................................................................. 111

 Clubiona maritima L. Koch, 1867 ......................................................................... 112

 Clubiona moesta Banks, 1896 .............................................................................. 112

 Clubiona norvegica Strand, 1900 ......................................................................... 112

 Clubiona obesa Hentz, 1847 ................................................................................ 113

 Clubiona pygmaea Banks, 1892 ........................................................................... 113

 Clubiona riparia L. Koch, 1866 ............................................................................. 113

 Clubiona trivialis C.L. Koch, 1843 ......................................................................... 114

 Family Corinnidae ...................................................................................................... 115

 Castianeira cingulata (C.L. Koch, 1841) ............................................................... 115

 Castianeira descripta (Hentz, 1847) ..................................................................... 116

 Castianeira gertschi Kaston, 1945 ........................................................................ 116

 Castianeira longipalpa (Hentz, 1847) ................................................................... 116

 Phrurotimpus alarius (Hentz, 1847) ...................................................................... 117

 Phrurotimpus borealis (Emerton, 1911) ................................................................ 117

 Scotinella divesta (Gertsch, 1941) ........................................................................ 118

 Scotinella pugnata (Emerton, 1890) ..................................................................... 118

 Family Gnaphosidae .................................................................................................. 119

 Callilepis pluto Banks, 1896 ................................................................................. 119

 Drassodes neglectus (Keyserling, 1887) ..............................................................120

 Drassyllus niger (Banks, 1896) .............................................................................120

 Drassyllus socius Chamberlin, 1922 ....................................................................120

 Gnaphosa muscorum (L. Koch, 1866) ..................................................................121

 Gnaphosa parvula Banks, 1896 ...........................................................................121

 Haplodrassus bicornis (Emerton, 1909) ...............................................................122

 Haplodrassus hiemalis (Emerton, 1909) ..............................................................122

 Haplodrassus signifer (C. L. Koch, 1839) .............................................................122

 Herpyllus ecclesiasticus Hentz, 1832 ...................................................................123

 Micaria elizabethae Gertsch, 1942 .......................................................................123
 Micaria gertschi Barrows & Ivie, 1942 ..................................................................123

 Micaria pulicaria (Sundevall, 1831) ......................................................................124

 Sergiolus ocellatus (Walckenaer, 1837) ...............................................................124

 Zelotes exiguoides Platnick & Shadab, 1983 .......................................................124

 Zelotes fratris Chamberlin, 1920 ..........................................................................125

 Zelotes hentzi Barrows, 1945 ...............................................................................125

 Family Philodromidae ................................................................................................127

 Philodromus cespitum (Walckenaer, 1802) ..........................................................127

 Philodromus exilis Banks, 1892 ............................................................................127

 Philodromus imbecillus Keyserling, 1880 .............................................................128

 Philodromus peninsulanus Gertsch, 1934 ............................................................128

 Philodromus placidus Banks, 1892 ......................................................................129

 Philodromus praelustris Keyserling, 1880 ............................................................129

 Philodromus rufus vibrans Dondale, 1964 ............................................................129

 Philodromus vulgaris (Hentz, 1847) .....................................................................130

 Thanatus formicinus (Clerck, 1757) ......................................................................130

 Thanatus rubicellus Mello-Leitão, 1929 ................................................................131

 Thanatus striatus C. L. Koch, 1845 ......................................................................131

 Tibellus maritimus (Menge, 1875) ........................................................................131

 Tibellus oblongus (Walckenaer, 1802) ..................................................................132

 Family Thomisidae ....................................................................................................133

 Coriarachne utahensis (Gertsch, 1932) ................................................................133

 Misumena vatia (Clerck, 1757) .............................................................................134

 Misumenops asperatus (Hentz, 1847) ..................................................................134

 Misumenops carletonicus Dondale & Redner, 1976 .............................................135

 Ozyptila distans Dondale & Redner, 1975 ............................................................135

 Ozyptila sp. (near distans Dondale & Redner, 1975) ...........................................136

 Tmarus angulatus (Walckenaer, 1837) .................................................................136

 Xysticus alboniger Turnbull et al., 1965 ................................................................136

 Xysticus ampullatus Turnbull et al., 1965 .............................................................137

 Xysticus chippewa Gertsch, 1953 ........................................................................137

 Xysticus elegans Keyserling, 1880 .......................................................................138

 Xysticus ellipticus Turnbull et al., 1965 .................................................................138

 Xysticus emertoni Keyserling, 1880 .....................................................................139

 Xysticus ferox (Hentz, 1847) ................................................................................139

 Xysticus luctans (C. L. Koch, 1845) ......................................................................140

 Xysticus luctuosus (Blackwall, 1836) ....................................................................140

 Xysticus punctatus Keyserling 1880 .....................................................................140

 Xysticus triguttatus Keyserling, 1880 ....................................................................141

 Xysticus winnipegensis Turnbull et al., 1965 ........................................................141
 Family Salticidae .......................................................................................................142

 Eris militaris (Hentz, 1845) ...................................................................................142

 Euophrys monadnock Emerton, 1891 ..................................................................144

 Evarcha hoyi (Peckham & Peckham, 1883) .........................................................144

 Ghelna canadensis (Banks, 1897) .......................................................................145

 Habronattus borealis (Banks, 1895) .....................................................................146

 Habronattus calcaratus maddisoni Griswold, 1987 ..............................................146

 Habronattus decorus (Blackwall, 1846) ................................................................146

 Habronattus viridipes (Hentz, 1846) .....................................................................147

 Marpissa grata (Gertsch, 1936) ............................................................................147

 Neon nelli Peckham & Peckham, 1888 ................................................................147

 Pelegrina fl aviceps (Kaston 1973) ........................................................................148

 Pelegrina fl avipes (Peckham & Peckham, 1888) .................................................148

 Pelegrina insignis (Banks, 1892) ..........................................................................149

 Pelegrina proterva (Walckenaer, 1837) ................................................................150

 Phidippus clarus Keyserling, 1885 .......................................................................150

 Phidippus purpuratus Keyserling, 1885 ................................................................151

 Phidippus whitmani Peckham & Peckham, 1909 .................................................151

 Salticus scenicus (Clerck, 1757) ..........................................................................152

 Sibianor aemulus (Gertsch, 1934) ........................................................................152

 Sitticus fl oricola palustris (Peckham & Peckham, 1883) ......................................152

 Sitticus pubescens (Fabricius, 1775) ....................................................................153

 Sitticus striatus Emerton, 1911 .............................................................................154

 Synageles noxiosus (Hentz, 1850) .......................................................................154

 Talavera minuta (Banks, 1895) .............................................................................154

 Tutelina similis (Banks, 1895) ...............................................................................155

Faunal Attributes and Habitat Associations .............................................................156

 Spider Taxa ................................................................................................................156

 Spider Numbers .........................................................................................................156

 Spider Foraging Guilds ..............................................................................................157

 Spider Sex Ratios ......................................................................................................158

 Collection Method Effects ..........................................................................................160

 Spider Habitat Associations .......................................................................................163

 Seashores ............................................................................................................163

 Seashore-backshores ...........................................................................................164

 Amelanchier-Rubus stand ....................................................................................164

 Old fi elds ...............................................................................................................164

 Old fi eld edges ......................................................................................................165

 Old fi eld-alder edge ..............................................................................................165

 Heaths ..................................................................................................................165

 Saltmarshes ..........................................................................................................166
 Saltmarsh edges ...................................................................................................166

 Salt meadow .........................................................................................................167

 Brackish marshes .................................................................................................167

 Freshwater marshes .............................................................................................168

 Freshwater marsh edges ......................................................................................168

 Freshwater pond ...................................................................................................169

 Freshwater streamsides .......................................................................................169

 Sphagnum bogs ...................................................................................................170

 Roadside disturbed areas .....................................................................................170

 Betula-Acer stand .................................................................................................170

 Bigtooth aspen stand ............................................................................................171

 Aspen-maple stand ...............................................................................................171

 White birch stand ..................................................................................................171

 Red maple-white birch stand ................................................................................171

 Coastal red spruce stands ....................................................................................171

 Mixed conifer stands .............................................................................................172

 Mixed conifer-hardwood stands ............................................................................172

 Mixed hardwood-conifer stands ............................................................................173

 Gravel pit ..............................................................................................................173

 School playground-disturbed area ........................................................................173

 Domestic habitats .................................................................................................174

 Invasive Species ........................................................................................................174

 Species-Range Extensions .......................................................................................175

 Faunal Similarities .....................................................................................................176

Estimated Faunal Richness ........................................................................................177

 Estimated Milbridge Fauna ........................................................................................177

Spider-Habitat Conservation ......................................................................................177

 Spider Conservation ..................................................................................................177

Acknowledgments .......................................................................................................186

Literature Cited ............................................................................................................187

Appendix ......................................................................................................................203
1

INTRODUCTION

Spiders are among the dominant predators in terrestrial ecosystems 

(Gertsch 1979, Ubick et al. 2005) that feed chiefl y on insects (Nyffeler et 

al. 1994, Riechert and Harp 1987). In turn, spiders are eaten by diverse 

wildlife, including other spiders, predatory wasps, fi sh, amphibians, reptiles, 

birds, and mammals (Foelix 1996, Gertsch 1979, Wise 1993). As obligate 

insectivores, spiders play signifi cant regulatory roles in forest ecosystems 

and agroecosystems (see reviews by Maloney et al. 2003, Nyffeler and 

Benz 1987, Nyffeler et al. 1994, Riechert and Bishop 1990, Riechert and 

Lockley 1984).

Spiders are found in virtually all terrestrial habitats (Turnbull 1973), 

including some freshwater and marine habitats, and on all continents 

except Antarctica (Cushing 2005). Globally, spiders are the seventh most 

diverse order of animals in terms of named species (Coddington and Levi 

1991). Some 108 families, 3,677 genera, and 39,725 species have been 

described to date (Platnick 2007), and estimates of extant species range as 

high as 170,000 species (Coddington and Levi 1991). However, despite 

their ubiquitous occurrence, relative abundance, and ecological importance, 

spiders are seldom included among the organisms collected, counted, and 

identifi ed during faunal surveys in Maine or elsewhere. In reference to 

faunal surveys, Grimaldi and Engel (2005, p. 12) state: “Another surprise 

about these kinds of studies is that there have been very few surveys of the 

insects or terrestrial arthropods of natural areas...Even though that type of 

study is so important to estimating how many species...exist.”

Here we describe our efforts to inventory the spiders associated with 

diverse communities and habitats of Milbridge, a minor civil division 

(MCD) encompassing some 6,289.8 ha of Washington County, ME. These 

efforts began in 1991 and continued through 2005. Our goals were to 1) 

visit and collect spiders in as many representative habitats of Milbridge 

as possible; 2) use a variety of collecting and sampling techniques (e.g., 

beating sheets, sweep nets, pitfall traps) to sample ground, herb, shrub, and 

tree strata; 3) identify the collected specimens to family, genus, and species 

where possible; and 4) compare and contrast the observed spider fauna of 

Milbridge with that found on Mount Desert Island in Hancock County, 

ME. Before the Milbridge study, Mount Desert Island was the area most 

comprehensively inventoried for spiders in Maine; an inventory conducted 

by William Procter and associates during the 1920s to1940s (Procter 1933, 

1938, 1946).

DANIEL T. JENNINGS, research 

entomologist (ret.), USDA, Forest 

Service, Northern Research Station, 

686 Government Road, Bradley, 

ME 04411. Corresponding author. 

Current address: P. O. Box 130, 

Garland, ME 04939

 Daniel.Jennings@umit.maine.edu

FRANK GRAHAM, JR., fi eld editor, 

National Audubon Society, 700 

Broadway, New York, NY 10003
2

STUDY SITES AND HABITATS

Study Sites

During the 15-year study period (1991-2005), about 60 sites within the MCD of Milbridge were 

sampled for spiders (Table 1, Figure 1). All study sites and collecting localities were within the East 

Coastal BioPhysical Region as defi ned by McMahon (1990). This region is characterized by low 

ridges surrounded by relatively fl at terrain; elevations are generally less than 100 ft (30.5 m); bedrock 

is predominantly igneous, with headlands and islands composed of biotite or granite (McMahon 

1990). The regional climate is characterized by cool summers and high annual precipitation that 

occurs chiefl y in the winter. Fog is common during the summer. Soils range from poorly developed, 

shallow acidic soils on ridgetops and outer peninsulas, to deep marine clays and glaciolacustrine 

deposits in low-lying areas, to glacial till and deep, moderately well drained loamy soils of ridge slopes 

(McMahon 1990). Prominent physical features in Milbridge include the Narraguagus River, which 

drains into Narraguagus Bay; Mill River, which drains into Flat Bay; Beaver Meadow Brook, which 

drains into Back Bay; the Gulf of Maine; fresh, brackish, and saltwater marshes; several offshore 

islands; and coastal granite ledges along the shorelines of bays, peninsulas, and some offshore islands 

(USGS topographic maps).

Two terrestrial ecosystems make up the Eastern Coastal Region: coastal spruce-fi r forests and coastal 

raised peatlands (McMahon 1990). The coastal spruce-fi r forests are comprised chiefl y of red spruce 

(Picea rubens Sarg.) and balsam fi r (Abies balsamea (L.) Mill.). In cutover and disturbed areas of these 

forests, prominent tree species include paper birch (Betula papyrifera Marsh.), red maple (Acer rubrum 

L.), and bigtooth aspen (Populus grandidentata Michx.).

Habitats

Specifi c habitats included seashores and associated cobble, sand and shingle beaches; seashorebackshores above wrack lines; granite ledges of seashores, islands, and headlands; freshwater marshes, 

brackish marshes, and saltmarshes; a salt meadow; riparian edges of rivers, streams, and a dry 

streambed; freshwater pond and beaver fl owages; herbaceous-shrub-small tree vegetation of old fi elds, 

old fi eld edges; disturbed roadsides and a school playground; a gravel pit; a Kalmia-Vaccinium heath; 

an Amelanchier-Rubus stand, an alder-old fi eld edge; mixed deciduous-coniferous woodland; inland 

ledges; deciduous forest stands of bigtooth aspen, paper birch, and mixed red maple-paper (white) 

birch; and coniferous forest stands of coastal red spruce, and mixed red spruce-balsam fi r. Special 

efforts were made to record specifi c spider-plant habitat associations; e.g., on common goldenrod

infl orescence; in silk retreat on curled red maple leaf; beating white spruce foliage. A list of common 

and Latin names of plants with associated spiders is found in the Appendix. The plant names given by 

Haines and Vining (1998) served as the primary source for both common and Latin names.

MATERIALS AND METHODS

Spider Collection Methods

Pitfall Traps

For terrestrial habitats, we used a modifi ed version of the large-capacity pitfall trap described by 

Houseweart et al. (1979). The new, modifi ed design consisted of three basic components: 1) a trap 

apron (17.8 x 17.8 cm) cut from 0.55-cm tempered hardboard, with a centrally located 9.9-cmdiameter hole whose circumference was routed to form a ledge (~ 0.3 cm deep x ~ 0.3 cm wide); 
3

Figure 1.—Distribution of spider collecting sites in Milbridge, a minor civil division of the East Coastal 

BioPhysical Region, Washington County, Maine.

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Addison

Harrington

Milbridge

Steuben

M24 M34

M04

M33

M12

M13

M18

M19

M21

M22

M23

M15

M25

M27

M28

M46

M31

M35

M37

M38 M53

M01

M40

M41

M51

M11

M30

M44

M50

M07

M06

M20

M43 M02

M05

M03

M42

M09

M32

M39

M52

M45

M55 M47

M17

M29

M14

M36

M57

M08

M58

M49

M16

M10

M48

M56

M54

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Maine

N

Milbridge

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4

2) a 473-ml plastic Solo® cup, fi lled to ~ 2.5 cm with 110 ml of propylene glycol (RV antifreeze) as 

a killing-preservative agent; and 3) a rain cover (22.9 x 22.9 cm) fashioned from ~ 0.32-cm exterior 

plywood, and supported by four 16d galvanized nails. The circumferential ledge suspends and holds 

the trap cup, whose lip fi ts fl ush with the surrounding apron. In Minnesota, Cutler et al. (1975) 

found that similar pitfall traps with aprons caught twice as many spiders as traps without aprons. The 

trap aprons also help to exclude soil and debris, thus yielding cleaner catches that are easier to sort.

Depending on available continuous microhabitat (i.e., same biotic-physical features), we deployed 

pitfall traps either in line transects or in 3 x 3 m grids. For both deployment designs, individual traps 

were spaced 3 m apart. Numbers of traps per site ranged from 5 to 9; periods of trapping ranged 

from May to September, but were not consistent among study years. However, trapping periods were 

always consistent among study sites for individual years.

After deployment, pitfall-trap contents were collected at about weekly intervals (range 5-14 

days); content collections were always consistent among study sites for individual years. For these 

collections, the trap cups were removed from the apron; their captured contents were passed through 

a small-mesh (kitchen) strainer and stored in plastic urine specimen jars (120 ml) containing 70-75 

percent isopropyl or ethyl alcohol. A pre-printed waterproof label was added to each jar; recorded 

data included collection locality, collection date(s) (i.e., trapping period), habitat, pitfall number, and 

collector’s name.

For aquatic habitats, we used fl oating pitfall traps similar to those described by Graham et al. (2003). 

The Milbridge traps were fashioned from 3.8-cm StyrofoamTM cut into 17 x 17 cm squares to form 

an apron or platform. A centrally located hole (~ 10 cm diameter) with a circumferential ledge similar 

to that described above was then scribed and cut with a razor blade. The ledge suspended and held 

a 473-ml plastic Solo® cup that contained ~ 140 gm of plaster of paris, which helped stabilize the 

fl oating platform. After the plaster of paris dried, a size 10 screw-eye (3.6 cm long) was screwed into 

the bottom of each cup. During deployment about 110 ml of propylene glycol (RV antifreeze) was 

added to each platform cup as a killing-preservative agent.

Unlike Graham et al. (2003), who tied their fl oating pitfall traps to an anchoring pole, we fashioned 

anchors from 149-ml plastic juice cups containing ~ 140-155 gm of quick-setting cement. Before the 

cement dried, a size 10 screw-eye was placed in the uppermost surface of the cement. Depending on 

water depth, variable lengths of nylon cord were attached to the screw-eye--one end to the platform 

cup, the opposite end to the anchoring cup. Such an arrangement eliminated the need for anchoring 

poles and allowed some horizontal movement of the fl oating trap on the water’s surface, depending 

on wind and currents.

To deploy these aquatic pitfall traps, we used a long (~ 2 m) pole to extend the anchoring cord and 

cup anchor out over the water 1-2 m from the bank. As the anchor was lowered into the water, 

the trap platform was released, thereby pulling the fl oating platform away from the bank. Slack in 

the anchor’s cord allowed some horizontal movement of the fl oating trap, depending on wind and 

currents. Five aquatic pitfall traps were deployed in each of four sampling sites: two beaver fl owages, 

one freshwater marsh pond, and one human-made pond.
5

To retrieve aquatic pitfall traps, we used a pole mounted with a shepherd’s hook to snag the trap’s 

anchoring cord. The fl oating trap and anchor were then pulled slowly to the bank. To retrieve trap 

contents (if any) and processing samples, we followed the same procedures as described above for 

terrestrial pitfall traps.

Sweep Nets

Standard insect-collecting nets were used to collect spiders from herbaceous-shrub-small tree 

vegetation. Two commonly used nets and sizes were BioQuip® Professional Series Insect Net, 30.5-

cm-diameter net of polyester or muslin netting, with a 91.5-cm handle; and BioQuip® Heavy Duty 

Sweep Net, 38-cm-diameter net of sailcloth with a 61-cm handle. Captured spiders were removed 

from sweep nets by aspirator or vial and stored in small jars or vials containing 70-75 percent 

isopropyl or ethyl alcohol. A penciled waterproof label bearing collection data was added to each jar 

or vial. Sweep nets were used chiefl y in old fi elds and in marshes, along edge habitats (e.g., roadsides, 

banks or shores of waterways, forest-woodland edges), and in forest openings and where understory 

vegetation was present. Other than recording numbers and frequency of captures by species, no 

attempts were made to quantify sweep-net collections either by volume or vegetation area swept.

Beating Sheets

To collect arboreal spiders from woody shrubs and trees, we used two styles and sizes of beating 

sheets: BioQuip® Canvas Beating Sheet, 71 x 71 cm square, with marine canvas (tan) supported by 

pivotal oak cross members; and BioQuip® Ripstop Beating Sheet, 1 x 1 m square, with lightweight 

nylon ripstock supported by four 71-cm sections of PVC tubing. Dislodged spiders were captured by 

aspirator or dry vial and then transferred to vials or jars containing 70-75 percent isopropyl or ethyl 

alcohol. A waterproof label bearing collection data was added to each storage container.

Tree-Bole Brush and Net

Tree boles were brushed with a medium soft handbrush (whitewash brush) and dislodged spiders 

were captured with a specially designed net (Dunn and Reeves 1980). The net was fashioned from a 

standard insect net. One-half of the net’s supporting metal rim was replaced with an elastic band that 

allowed close placement of the net against a tree bole. Dislodged spiders were retrieved by aspirator or 

dry vial and transferred to vials or jars containing alcohol. A waterproof label bearing collection data 

was added to each storage container.

Litter Condenser-Extraction

Samples of litter were collected from forest fl oors and marsh edges and condensed by a two-handled 

litter condenser, similar to that illustrated in the latest BioQuip® Products (2007), online catalog. 

However, the litter condenser used at Milbridge is equipped with a single 6 x 6 mm mesh screen, 

instead of two screens (mesh sizes 1 cm and 50 mm). Condensed litter was either 1) sorted by hand 

in white plastic or porcelain trays, or on black plastic sheets; or 2) processed by modifi ed Berlese 

funnels. Resultant specimens were processed as described earlier.

Searches

Spiders were also collected by searching under rocks, logs, ground debris, and loose dead bark of 

trees and snags. Numerous specimens of several species were taken by examining interior and exterior 

walls, fl oors, and ceilings of houses, sheds, cottages, and other buildings. In such synanthropic 
6

habitats, spiders were captured by aspirator, dry vial, or dry jar; specimens were processed and stored 

like the other collections.

Collection Site and Habitat Descriptors

Global Positioning System Coordinates

At each sampling-collection site, latitude-longitude coordinates were determined by GarminTM 

GPS 38 and GarminTM GPS map 60CS hand-held meters. The coordinates were then plotted using 

Maptech® Terrain Navigator 2002 and GIS Arcview® 9.1 software in WMF format, Region 601, for 

preparation of Figure 1.

Habitat Descriptors

Habitat descriptions were based chiefl y on dominant and co-dominant vegetation type (e.g., BetulaAcer stand, mixed hardwood-conifer, mixed conifer-hardwood) and on general physiographic features 

(e.g., seashore, seashore-backshore, gravel pit).

Specimen Identifi cations and Museum Deposits

Spider Identifi cations

Collected specimens were processed in the laboratory and examined with stereo-zoom microscopes 

SwiftTM MZ 800, 10.5-67.5x magnifi cation; LeicaTM MZ-8, 10.1-80.0x magnifi cation), both 

equipped with fi ber-optic lighting. With few exceptions, both authors examined and identifi ed 

each collected specimen. Most adult spiders were identifi ed to species following Kaston (1981) and 

other consulted sources, which included Opell and Beatty (1976) for the Hahniidae, Leech (1972) 

for the Amaurobiidae, Chamberlin and Gertsch (1958) for the Dictynidae, Dondale and Redner 

(1982) for the Clubionidae, Dondale and Redner (1978b) for the Philodromidae and Thomisidae, 

Dondale and Redner (1990) for the Lycosidae and Pisauridae, and Platnick and Dondale (1992) 

for the Gnaphosidae. For determining species of Linyphiidae (Linyphiinae, Erigoninae), numerous 

taxonomic papers were consulted; a few diffi cult specimens were sent to colleagues (Drs. C. D. 

Dondale, Ottawa, and P. J. van Helsdingen, Leiden, The Netherlands) for identifying or verifying 

species. Arrangement and enumeration of spider taxa generally follow that of Platnick (2007), except 

families are listed by foraging guild.

Museum Specimen Deposits

With few exceptions, representative specimens of all identifi ed species are slated for deposit in the 

arachnid collections of the Museum of Comparative Zoology at Harvard University. Females of two 

unidentifi ed erigonids (Tapinocyba sp., and Unknown genus, species), plus one unidentifi ed male 

Ozyptila, are in the Canadian National Collection at Ottawa. Males of a new Pirata were sent to 

James C. Cokendolpher for species description.

Data Summaries and Analyses

Data Summaries

Specimen collection data (i.e., locality, date collected, method, habitat, and collector) were entered 

by species into versions 1.61 and 2.02 of Biota©: The Biodiversity Database Manager for tabulation 

and summarization. Only adult spiders were included in the database. All 12 of Biota’s core tables 

were used with individual specimens receiving the following codes: specimen, species, collection, and 

locality. For each species, summary tables were printed showing the number of specimens collected by 

locality, date(s), collection method, specimen sex, habitat, and microhabitat.
7

Data Analyses

For most variables, descriptive statistics were used to calculate means and standard errors. 

Nonparametric procedures (Sokal and Rohlf 1981) were used to compare observed vs. expected 

proportions at P = 0.05.

Sørensen’s similarity quotient (QS) as defi ned by Price (1975) was used as a qualitative measure of 

similarity between the Milbridge spider fauna and that of Mount Desert Island (MDI). The latter 

was inventoried chiefl y by William Procter and associates during the 1920s-1940s (Procter 1933, 

1938, 1946). The formula used was QS = 2c x 100/(a + b), where a = the number of spider taxa 

(family, genera, species) in study A (Milbridge); b = the number of spider taxa in study B (MDI); 

and c = the number of spider taxa common to both studies. Only identifi ed taxa were included in 

these comparisons; unknown species and species represented by juveniles were excluded. Quantitative 

indices of similarity (e.g., Bray-Curtis, Simpson’s) were not appropriate because spider abundances 

were not always recorded by species for the MDI fauna.

However, before such faunal comparisons could be made, recent species synonymies and other 

taxonomic changes (e.g., reassignment of genera, species to other families) (Platnick 2007) had to be 

considered. For example, Procter (1946, p. 525) listed Phrurotimpus palustris Banks as a member of 

the Clubionidae. This species has since been designated as a junior synonym of Phrurotimpus alarius 

(Hentz 1847), and subsequently the genus Phrurotimpus was reassigned fi rst to the Gnaphosidae 

(Brignoli (1983), then to the Liocranidae (Platnick 1989, 1993, 1997), and most recently to the 

Corinnidae (Bosselaers and Jocqué 2002, Platnick 2007). Similar revisions were made for other 

outdated taxa listed by Procter (1946) for the Mount Desert Island fauna.
8

Site # Locality name 1 Coordinates 2 Habitats sampled

M01 Graham’s property, 170 Wyman Rd. 44° 31.6′ N, -67° 52.1′ W Seashore; Seashore-Backshore; Mixed DeciduousConiferous Woodland; Amelanchier-Rubus Stand; 

Betula-Acer Stand; Freshwater Pond Edge; Old Field; 

Old Field-Edges; Old Field-Alder Edge; Arboreal; 

Domestic (house, screen house, shed, cabin, boats, etc.)

M02 Rays Point Road, roadside parking 

1.6 km E jct. Back Bay Road & 

Ray’s Point Road

44° 33.8′ N, -67° 50.2′ W Roadside Vegetation, Mixed Conifer 

M03 Bois Bubert Island 44° 26.5′ N, -67° 52.3′ W Freshwater Pond Edge

M04 McClellan Park, Narraguagus Bay 44° 29.2′ N, -67° 51.4′ W Coastal Red Spruce Forest 

.......................................... 44° 29.1′ N, -67° 51.3′ W Wet Meadow 

.......................................... 44° 29.2′ N, -67° 51.3′ W Roadside, Coastal Red Spruce Forest

M05 Kansas Road, Narraguagus River

0.5 km NW of jct. Kansas Road &

Kennedy Hwy. (Rt. 1A).

44° 33.2′ N, -67° 53.4′ W Old Field 

......................................... 44° 33.1′ N, -67° 53.4′ W Old Field-Brackish Marsh Edge

M06 Beaver Meadow Brook, Back Bay

Road

44° 33.3′ N, -67° 52.1′ W Saltmarsh; Saltmarsh Edge; Arboreal 

M07 Ficketts Point Road, Cemetery

1.3 km SE jct. Ficketts Point Road

& Bayview Street

44° 32.2′ N, -67° 51.7′ W Kalmia-Vaccinium Heath 

M08 Tom Leighton Point 44° 27.9′ N, -67° 52.1′ W Seashore; Seashore-Backshore

M09 Bois Bubert Island 44° 27.1′ N, -67° 52.3′ W Old Field-Abandoned Farm; Arboreal

M10 Hubbell property, 0.2 km S jct.

Wyman & Tom Leighton Pt. Roads

44° 30.4′ N, -67° 51.5′ W Mixed Conifer-Hardwood; Ledges; Arboreal

M11 Gravel pit, 2.4 km NW of Milbridge 44° 33.1′ N, -67° 54.7′ W Gravel Pit 

M12 Petit Manan NWR, 0.6 km SW jct.

Wyman & Marsh Roads

44° 30.3′ N, -67° 52.0′ W Freshwater Marsh Pond 

M13 Heath Road and Mill River 44° 36.0′ N, -67° 52.2′ W Mixed Hardwood-Conifer; Roadside (culvert); 

Riparian-Streamside; Arboreal

M14 Wyman, Trail to Sawyer Marsh

0.3 km SW jct. Wyman & Marsh Roads

44° 30.4′ N, -67° 51.8′ W Mixed Hardwood-Conifer (understory); Sand Pit

M15 Wyman, Sawyer Saltmarsh, 1.4 km

SW jct. Wyman & Marsh Roads

44° 29.8′ N, -67° 52.2′ W Saltmarsh 

M16 McClellan Park, Narraguagus Bay 44° 29.1′ N, -67° 51.2′ W Seashore Ledges

M17 Cain’s Creek N of Graham’s 

property, W of Timmy Point

44° 31.7′ N, -67° 52.4′ W Saltmarsh

M18 Rays Point Road, Back Bay Road

1.1 km NW jct. Ray’s Point & Back

Bay Roads

44° 33.8′ N, -67° 50.6′ W Roadside (culvert); Arboreal 

M19 Heath Road, 1.1 km NW jct. Heath

& Kennedy Hwy., (Rt. 1A)

44° 35.4′ N, -67° 52.2′ W Mixed Hardwood-Conifer; Arboreal

M20 Bayside Supermarket, 23 Main Street 44° 33.2′ N, -67° 52.9′ W Domestic (paved parking lot)

M21 Headquarters, Petit Manan NWR,

14 Water Street

44° 32.3′ N, -67° 52.8′ W Domestic (building)

Table 1.—Sampling sites, coordinates, and habitats of spiders collected in Milbridge, Washington County, ME, 1991-2005

continued
9

M22 Petit Manan NWR, Sawyer Division,

1.2 km SW jct. Wyman & Marsh Roads

44° 29.9′ N, -67° 52.7′ W Bigtooth Aspen Stand

M23 Bois Bubert Island 44° 27.4′ N, -67° 52.2′ W Seashore; Seashore-Backshore; Sphagnum Bog; 

Mixed Conifer (opening)

M24 Petit Manan NWR, Sawyer Division,

0.8 km SW of Wyman & Marsh Roads

44° 30.4′ N, -67° 52.2′ W Red Maple Sapling Stand; Red Maple Sapling 

(understory)

M25 Petit Manan NWR, Sawyer Division,

1.1 km SW of Wyman & Marsh Roads

44° 30.0′ N, -67° 52.1′ W White Birch Stand; White Birch (understory) 

M26 Petit Manan NWR, Sawyer Division,

1.4 km SW of Wyman & Marsh Roads

44° 29.9′ N, -67° 52.2′ W Saltmarsh Edge

M27 Petit Manan NWR, Sawyer Division

1.6 km NE of jct. Joe Leighton & Bar

Island Roads

44° 29.8′ N, -67° 52.3′ W Red Maple-White Birch Stand

M28 E. Milbridge, Rt. 1A & Mill River 44° 35.2′ N, -67° 51.7′ W Roadside Disturbed Area 

M29 W. side of 170 Wyman Road 

1.1 km SE jct. Wyman Road 

& Rt. 1 (Main Street)

44° 31.5′ N, -67° 52.3′ W Roadside; Arboreal; Domestic (mailbox)

M30 Freeman’s Marsh (roadside), 1.6 km

SW jct. Washington Street & Rt. 1

44° 31.2′ N, -67° 53.8′ W Roadside; Roadside-Alder Edge; Mixed Conifer

(understory)

M31 Old County Road, 0.7 km W jct. 

Heath & Lynch Hill Roads

44° 35.9′ N, -67° 53.0′ W White Pine-Mixed Conifer

M32 Heath Road, 1.3 km NW of Rt.1A 44° 35.4′ N, -67° 53.0′ W Roadside; Mixed Conifer-Hardwood

M33 Freeman’s Marsh (meadow), 1.6 km

SW jct. Washington Street & Rt. 1

44° 31.2′ N, -67° 53.8′ W Freshwater Marsh 

M34 Petit Manan NWR, 0.5 km SW of 44° 30.4′ N, -67° 51.9′ W Riparian (dry streambed); Mixed Hardwood-Conifer; 

Wyman & Marsh Roads Ledge

M35 Kansas Road, 1.1 km NW jct. Kansas 

Road & Kennedy Hwy., (Rt. 1A).

44° 33.5′ N, -67° 53.6′ W Brackish Marsh

M36 Petit Manan NWR, 0.7 km SW jct.

Wyman & Marsh Roads; trail NE

of freshwater marsh pond

44° 30.2′ N, -67° 52.0′ W Mixed Conifer-Hardwood (understory)

M37 Red Barn Motel property, 3 North

Main Street

44° 32.4′ N, -67° 53.0′ W Old Field; Old Field Edge

M38 Flaherty Road, E. Milbridge, 0.3 km

S jct. Flaherty & Kennedy Hwy., (Rt. 1A)

44° 34.8′ N, -67° 51.8′ W Bog (Sphagnum-cranberry-rush)

M39 Flaherty Road, E. Milbridge, 0.3 km

S jct. Flaherty Road & Kennedy Hwy., 

(Rt. 1A)

44° 34.7′ N, -67° 51.7′ W Red Maple (understory); Red Maple-Red Oak 

(ledge)

M40 Old County Road, 1.1 km W jct. 

Heath & Lynch Hill Roads

44° 36.0′ N, -67° 53.3′ W Lowbush Blueberry Field

M41 Petit Manan NWR, Sawyer Saltmarsh,

1.3 km SW jct.Wyman & Marsh Roads

44° 30.0′ N, -67° 52.3′ W Saltmarsh Edge (impaled leaves) 

M42 Bois Bubert Island 44° 26.9′ N, -67° 52.4′ W Mixed Conifer Ledges

M44 Evergreen Cemetery, Rt. 1 44° 31.3′ N, -67° 53.7′ W Roadside-Domestic 

M45 Jct. Rt. 1 & Wyman Road 44° 31.9′ N, -67° 52.9′ W Roadside-Riparian Streamside

M46 Pinkham residence, 1.6 km SW jct.

Washington Street & Rt. 1, near

Freeman’s Meadow

44° 31.2′ N, -67° 53.9′ W Domestic 

Table 1.—continued

Site # Locality name 1 Coordinates 2 Habitats sampled

continued
10

M47 Milbridge, Washington Street, 0.2 km S. 

jct. Washington & Cottage Streets

44° 32.1′ N, -67° 53.1′ W Riparian-Freshwater Streamside

M48 Bobby Creek Causeway, N shore 44° 29.0′ N, -67° 52.6′ W Seashore-Backshore; Ledges

M49 S of McClellan Park, Narraguagus Bay 44° 29.1′ N, -67° 51.2′ W Seashore-Backshore

M50 Ray Point, Harrington Bay, 4.0 km

SE jct. Back Bay & Rays Point Roads

44° 32.5′ N, -67° 49.1′ W Seashore (cobble beach); Seashore-Backshore

M51 Old Fairgrounds, NW Milbridge

5.1 km NW jct. Kansas Road & 

Kennedy Hwy., (Rt. 1A)

44° 35.5′ N, -67° 54.7′ W Mixed Conifer (opening); Roadside

M52 Chapman residence, 88 Main Street 44° 32.0′ N, -67° 52.9′ W Domestic (house, barn)

M53 Flaherty Road, E. Milbridge, 0.3 km

SE jct. Flaherty Road & Kennedy 

Hwy., (Rt. 1A)

44° 34.8′ N, -67° 51.7′ W Mixed Hardwoods; Aspen-Maple

M54 Bar Island, McCormick property,

281 Bar Island Road

44° 28.4′ N, -67° 52.5′ W Mixed Conifer (litter); Arboreal

M55 Milbridge Elementary School,

39 Washington Street

44° 32.1′ N, -67° 53.3′ W Riparian-Beaver Flowage; School PlaygroundDisturbed Area

M56 Bar Island, near Bobby Creek

Causeway

44° 28.9′ N, -67° 52.6′ W Salt Meadow 

M57 N of Tom Leighton Point, parking

3.6 km S jct. Wyman & Tom Leighton

Point Roads

44° 28.6′ N, -67° 52.2′ W Brackish Marsh 

M58 S of McClellan Park, 2.8 km S jct.

Wyman & Tom Leighton Point Roads

44° 29.0′ N, -67° 51.2′ W Seashore Brackish Marsh 

1 Locality names, including roads, points, bays, streams, and marshes, are per The Maine Atlas and Gazetteer, 29th ed. 2006.

 Yarmouth, ME: DeLorme. 78 p.

2 Coordinates were determined by GarminTM GPS 38 and GarminTM GPSmap 60CS hand-held meters, and then plotted using 

Maptech® Terrain Navigator 2002 and GIS Arcview 3.2 software in WMF format, Region 601, for preparation of Map 1. 

Table 1.—continued

Site # Locality name 1 Coordinates 2 Habitats sampled
11

ANNOTATED FAUNAL LIST: INTRODUCTION

Spider families are grouped fi rst by two basic foraging guilds, the web spinner families and the hunter 

families. Within each guild, the families are listed according to the classifi cation of Platnick (2007). 

Families and genera of North American spiders can be identifi ed using the keys and descriptions in 

Ubick (2005). Species identifi cations require consultations with monographs and revisionary works; 

pertinent sources are listed for each family and species included herein. The spider identifi cation 

manuals of Dondale and Redner (1978b, 1982, 1990), Dondale et al. (2003), and Platnick and 

Dondale (1992) are especially helpful for identifying species found in New England and maritime 

Canada. Excellent illustrations of the spiders of Québec are provided by Paquin and Dupérré (2003). 

Although somewhat outdated, B. J. Kaston’s (1981) classical “Spiders of Connecticut” gives insights 

into spider natural history.

With modifi cation, our annotation style and format follows that of Burian and Gibbs (1991) 

“Mayfl ies of Maine: An Annotated Faunal List” and Bélanger and Hutchinson (1992) “Liste Annotée 

des Araignées (Araneae) du Québec.” Taxonomy includes literature citations of pertinent revisions, 

and sources of keys, descriptions, and illustrations. Collection records are listed by Milbridge 

collection-site number and include the number of males and females collected at each site. The 

collecting sites are defi ned in Table 1 and include location name, latitude-longitude coordinates, and 

habitats sampled. Map 1 shows the distribution of all sampled sites. Method(s) include the number 

of males and females taken by each sampling method; e.g., beating cloth, pitfall trap, search, sweep 

net. Month(s) include the cumulative number of males and females taken each month. Habitat 

associations include habitat (e.g., saltmarsh, freshwater marsh, old fi eld) and microhabitat descriptors 

(e.g., on cobble beach; on red spruce foliage; in marsh litter), followed by the number of male and 

female specimens per habitat. A few specimens were received bearing no information about method, 

date, or habitat; these are indicated by the standard abbreviation “n. d.” for no data. Species regional 

distributions include only the New England States (ME, NH, VT, MA, CT, RI, NY) and the 

Maritime Provinces of Canada (ON, QC, NB, NS, PE, NF, LB). State and provincial abbreviations 

follow standard postal codes. If additional geographic-distributional data are needed, they can be 

found elsewhere; e.g., species-generic revisions, identifi cation manuals, and some regional faunal 

lists. To avoid redundancy, geographic-citation sources are somewhat cumulative. The source(s) 

of the earliest known Maine record (if any) is listed fi rst, followed by the source(s) with the most 

comprehensive listing of species range (usually revisionary work), and lastly, by any recent additions. 

New State records are indicated by (*), new records for New England by (**), and new National 

records by (***).
12

ANNOTATED FAUNAL LIST: 

WEB SPINNER FAMILIES

FAMILY PHOLCIDAE

The pholcids or cellar spiders have long thin legs with fl exible tarsi. The anterior median eyes are the 

smallest or lacking, while the other eyes are arranged in two triads. The webs are either sheet-like or 

irregular, with the spiders hanging in an inverted position, either beneath or in the lower portions of 

the web. The webs are usually found in dark corners or ceilings of basements and cellars. After egglaying, the eggs are covered by only a few strands of silk and held by the female in her chelicerae. See 

Huber (2005) for additional information on family characteristics, notes on natural and taxonomic 

histories, and a key to the genera found in North America north of Mexico. Identifi cation keys for 

North American species are limited and, in some cases, misleading; however, see revisionary works by 

B. A. Huber as listed in Huber (2005) and Platnick (2007). Several species of Pholcus have yet to be 

described.

Members of the Pholcidae are more common in warmer regions; however, several species are 

cosmopolitan, especially those associated with domestic habitats. Thus far, only two genera and three 

species have been found in Maine; a single genus and species have been taken in Milbridge. Most 

likely, the cosmopolitan Pholcus phalangioides (Fuesslin 1775) also occurs in Milbridge, but remains 

to be found.

Pholcus manueli Gertsch, 1937

Taxonomy: Gertsch (1937); Kaston (1981, as Pholcus opilionoides (Schrank, 1781)); Senglet (2001).

Records: M01 (1 female); M21 (1 male). N = 2 adults; 1 male, 1 female.

Method: search (1 male, 1 female).

Months: September (1 male); November (1 female).

Habitats: in web, house cellar (1 female); by fl oor and wall, government building (1 male).

Regional Distribution: ME (Jennings, unpubl.); NY (Kaston 1981, as Pholcus opilionoides 

(Schrank)).

Note: Gertsch (1937) fi rst described Pholcus manueli based on specimens collected in a garage in 

Mendham, NJ. Later, Kaston (1977, 1981) indicated that Gertsch’s manueli was a junior synonym 

of Pholcus opilionoides (Schrank, 1781), a European species. However, Senglet (2001) challenged 

Kaston’s synonymy and provided evidence the two species were clearly separate. Before the Milbridge 

study, specimens of P. manueli were taken from a fi eld laboratory in Bradley, and from a building 

on the University of Maine campus in Orono, both localities in Penobscot County, ME. Like other 

pholcid spiders, P. manueli appears to be a synanthropic species closely associated with domestic 

habitats.
13

FAMILY THERIDIIDAE

Most theridiids or cobweb weavers have a ventral row of serrated bristles on tarsus IV that are used to 

comb silk from the spinnerets. Males of some species lack a comb. Spiders of this family spin irregular 

mesh webs, with the resident spider hanging in an inverted position in the web. Captured prey is 

enswathed with viscid silk by the hind legs. Members of this large family are found in diverse habitats 

depending on species and their habits; some are arboreal, living in and among foliage of conifers and 

hardwoods; others live in the herb-shrub layer, often folding and tying leaves to form egg retreats; still 

others are ground dwellers and are found in litter or under rocks.

This family is represented in Maine by 18 genera and 52 species, including some invasive species, 

and at least three medically important species of Latrodectus or widow spiders. None of the latter 

are known to occur in Milbridge. Thus far, only 15 genera and 27 species of theridiids have been 

found in Milbridge; no doubt other species can be found. Most of the North American genera 

of Theridiidae have been revised by H. W. Levi, Museum of Comparative Zoology, at Harvard 

University. These excellent revisionary works include descriptions and illustrations of species and 

diagnostic keys to genera and species. For listings of revisionary literature, see Levi and Randolph 

(1975) and Levi (2005).

Achaearanea globosa (Hentz, 1850)

Taxonomy: Levi (1955a, 1963a); Kaston (1981); Paquin and Dupérré (2003).

Records: M01 (1 female); M06 (1 female); M07 (1 female); M22 (1 female); M28 (1 female). N = 5 

adult females.

Methods: pitfall traps (2 females); search (1 female); sifted litter-hand sorted (1 female); sweep net 

(1 female).

Months: May (1 female); June-July (1 female); July (1 female); August (2 females).

Habitats: forest-fl oor litter, bigtooth aspen stand (1 female); litter, saltmarsh (1 female); sifted litter, 

Kalmia-Vaccinium heath (1 female); sweeping roadside vegetation, disturbed area (1 female); in web 

on lawn chair, old fi eld edge (1 female).

Regional Distribution: ME (Bryant 1908, as Theridion globosum Hentz); CT (Kaston 1981); NY 

(Crosby and Bishop 1928, as Theridion globosum Hentz); ON, QC (Levi and Randolph 1975).

Achaearanea tabulata Levi, 1980

Taxonomy: Levi (1980a); Paquin and Dupérré (2003).

Records: M01 (40 males, 62 females); M04 (1 female); M06 (4 females); M07 (1 female); M13 

(7 females); M16 (1 male, 3 females); M18 (3 females); M46 (4 females); M47 (3 females); M48 

(5 females). N = 134 adults; 41 males, 93 females.

Methods: pitfall traps (4 females); search (41 males, 88 females); n. d. (1 female).
14

Months: May (1 male, 4 females); June (16 males, 18 females); July (15 males, 31 females); August 

(8 males, 30 females); September (8 females); October (1 male, 2 females).

Habitats: litter, saltmarsh (3 females); litter, Kalmia-Vaccinium heath (1 female); descending on 

dragline from cinquefoil (1 male); in rolled leaf retreat on large rock, old fi eld (1 female); in rolled 

leaf inside cement block (1 female); on foliage, old fi eld (1 male); in webs, seashore ledges (1 male, 

8 females); in webs, roadside culvert, mixed conifer-low wet area (3 females); in webs, roadside 

culvert, mixed hardwood-conifer (6 females); in webs under bridge, freshwater stream (3 females); 

in web with Callobius bennetti prey, bridge guardrail, freshwater stream (1 female); near bridge, 

saltmarsh (1 female); in woodpile (2 males, 2 females); in retreats, outdoor woodbin (3 females); in 

web, woodbin (1 female); in vegetable garden (1 female); in web, compost drum, feeding on Tipulid 

(1 female); running outside compost bin (1 female); in webs, boat and canoe (1 male, 2 females); 

in webs, interior walls, window, fl oor, shoe, and door, house garage (5 males, 10 females); in webs, 

house basement or cellar (2 males, 6 females); on cellar stairs (1 female); in cellar (1 female); in webs, 

exterior (shingled) walls and window, house (1 male, 6 females); on exterior walls, house (10 males, 

6 females); in webs, exterior foundation, house (5 males); on exterior foundation, house (3 females); 

in webs, or on interior walls, ceiling, window, and screen door (4 males, 2 females); in bathtub, house 

(2 males); in bedroom, house (1 male); on house deck (1 male); dead among books, house deck (1 

male); in webs, inside or on shed (2 males, 11 females); on fl oor, screened house near freshwater pond 

(1 female); under lawn furniture (1 female); on back of lawn chair (1 female); in web, exterior wall, 

camp (cabin) near seashore (1 male); in webs under sun porch, house (3 females); in web under porch 

step with beetle prey (1 female); n. d. (1 female).

Regional Distribution: ME (Jennings, unpubl.), NY (Levi 1980a); ON, QC, NB (Dondale et al. 

1994).

Note: This is an invasive species fi rst discovered and described from a single female collected in 

1976 on Long Island, NY (Levi 1980a). It since appears to have spread inland and northward, and 

as far west as Chicago, IL (B. Cutler, pers. comm.). The country of origin is unknown. Levi (1980a) 

concluded that A. tabulata was not a European species; historically, the theridiid fauna of Europe has 

been well studied. More recently specimens have been found in urban areas of Germany and Austria 

(see Dondale et al. 1994, lit. cited). Dondale et al. (1994) indicated this introduced species might 

have originated in Japan, Korea, or some other southeastern Asian country or countries. Because of its 

close resemblance to Parasteatoda tepidariorum (C. L. Koch, 1841) and possible misidentifi cations, A. 

tabulata may have a wider distribution in New England and Canada than previously known.

Crustulina sticta (O. Pickard-Cambridge, 1861)

Taxonomy: Levi (1957a); Kaston (1981); Paquin and Dupérré (2003).

Records: M01 (1 male); M23 (1 male). N = 2 adult males.

Methods: search (2 males).

Months: May (1 male); July (1 male).
15

Habitats: [on] Spartina, rocky beach, seashore (1 male); on exterior wall, house (1 male).

Regional Distribution: ME (Procter 1946, as Crustulina borealis Banks); NY (Crosby and Bishop 

1928); NH, MA, CT, ON, QC, NS, NF (Levi 1957a). 

Dipoena nigra (Emerton, 1882)

Taxonomy: (Levi 1953); Kaston (1981); Paquin and Dupérré (2003).

Records: M06 (1 male, 6 females); M23 (1 male); M49 (2 females). N = 10 adults; 2 males, 8 

females.

Methods: beating cloth (1 male, 7 females); sweep net (1 male, 1 female).

Months: June (1 male); July (1 male, 8 females).

Habitats: beating white spruce foliage, saltmarsh edge (1 male, 5 females); beating red spruce foliage, 

seashore-backshore (2 females); sweeping vegetation, saltmarsh edge (1 female); sweeping Kalmia sp. 

near freshwater pond, mixed conifer (1 male).

Regional Distribution: ME (Emerton 1882, as Steatoda nigra Emerton); NY (Crosby and Bishop 

1928); CT (Kaston 1981); NH, VT, MA (Levi 1953); QC (Bélanger and Hutchinson 1992).

*Enoplognatha intrepida (Sørensen, 1898)

Taxonomy: Levi (1957b); Kaston (1981); Paquin and Dupérré (2003).

Record: M01 (1 female). N = 1 adult female.

Method: search (1 female).

Month: July (1 female).

Habitat: under white (paper) birch bark (1 female).

Regional Distribution: ME (this study); NH, MA, CT, NY, ON (Levi 1957b); QC Paquin and 

Dupérré (2003).

Enoplognatha ovata (Clerck, 1757)

Taxonomy: Levi (1957b); Kaston (1981, as Theridion redimitum (Linnaeus, 1758)); Paquin and 

Dupérré (2003).

Records: M01 (14 males, 75 females); M04 (1 female); M07 (2 males, 18 females); M14 (6 males, 

1 female); M17 (2 males, 5 females); M23 (1 male, 1 female); M28 (2 males, 12 females); M29 (1 

female); M30 (3 females); M47 (1 male, 5 females); M50 (2 males, 6 females). N = 158 adults; 30 

males, 128 females.
16

Methods: beating cloth (2 males, 13 females); pitfall traps (2 males, 18 females); search (5 males, 

23 females); sweep net (21 males, 74 females).

Months: May (2 males, 18 females); June (1 male); July (21 males, 73 females); August (6 males, 

29 females); September (8 females).

Habitats: beating spruce (Picea sp.) branches (1 female); beating foliage, alder (Alnus sp.) thicket 

(2 males, 10 females); beating understory red spruce foliage, mixed conifer-hardwoods (1 female); 

beating red spruce foliage, roadside (1 female); sweeping Vaccinium and Myrica, old fi eld (6 females); 

sweeping vegetation, old fi eld (1 male, 10 females); sweeping vegetation, old fi eld edge (3 males, 11 

females); sweeping Vaccinium and low shrubs, old fi eld edge (2 females); sweeping aster and raspberry 

(1 female); sweeping understory low shrubs, Betula-Acer woods (1 male, 5 females); sweeping 

meadowsweet stand (1 female); sweeping vegetation, freshwater pond edge (1 male, 2 females); 

sweeping grasses, seashore-backshore (1 male, 2 females); sweeping grasses and sedges, seashorebackshore (1 female); sweeping understory vegetation, spruce-mixed hardwood (1 female); sweeping 

grasses and forbs along trail, mixed hardwood-conifer (6 males, 1 female); sweeping grasses, saltmarsh 

(1 male); sweeping grasses and shrubs, saltmarsh (1 male, 4 females); sweeping vegetation, saltmarsh 

(1 female); sweeping Vaccinium sp., mixed conifer (1 male); sweeping roadside vegetation, disturbed 

land (5 females); sweeping roadside white and yellow clover, disturbed area, mixed hardwood-conifer 

(2 males, 7 females); sweeping roadside grasses, forbs, ferns, and rushes, alder-aspen edge (3 females); 

sweeping marsh vegetation, freshwater streamside, riparian (1 male, 5 females); sweeping grasses, 

forbs, and shrubs, seashore-backshore (2 males, 6 females); litter, Kalmia-Vaccinium heath (2 males, 

18 females); in rolled wild raspberry leaf (1 female); in bigtooth aspen leaf, freshwater pond edge 

(1 female); in web, garden-heliotrope fl ower head (1 female); in folded alder leaf (1 female); in web 

on chokecherry (1 female); in silken retreat, folded seaside goldenrod leaf (1 female); in rolled leaf, 

seaside goldenrod, seashore-backshore (1 female); among clump of seaside goldenrod, seashorebackshore (1 female); in webs, in or on alders, alder thicket (2 males, 5 females); inside folded leaf 

(n. d.) with egg sac (1 female); in rolled leaf of witch alder, ornamental garden (1 female); on white 

(paper) birch with prey (1 female); on marsh grass with prey, saltmarsh (1 female); descending silk 

line from tree (n. d.), woodland (1 male); on woodland fl oor (1 male); on small rocks, seashore (1 

female); in vegetable garden (2 females); on pea plant, vegetable garden (1 female); inside boat, old 

fi eld (1 female); on house deck (1 female); descending silk line from room ceiling, house (1 male).

Regional Distribution: ME (Procter 1946, as Theridion redimitum (Linnaeus)); MA, RI, NY, ON 

(Levi 1957b); CT (Kaston 1981); QC (Reillo 1989, Bélanger and Hutchinson 1992).

Note: This is an introduced species that probably originated in Europe where it is widely distributed 

(Levi 1957b). Early records of this species in North America are chiefl y from coastal regions of 

the Pacifi c Northwest and the Atlantic seaboard (Levi 1957b, Map 1); such distribution suggests 

a maritime introductory mode. More recently, populations of E. ovata have been detected further 

inland in New York, along the St. Lawrence River in Québec (Reillo 1989), and in inland regions of 

Québec (Bélanger and Hutchinson 1992).

Euryopis funebris (Hentz, 1850)

Taxonomy: Levi (1954a), as E. limbata (Walckenaer, 1841); Kaston (1981); Paquin and Dupérré 

(2003).
17

Records: M06 (1 female); M10 (1 female); M51 (1 female). N = 3 adult females.

Method: beating cloth (3 females).

Months: May (1 female); June (1 female); July (1 female).

Habitats: beating white spruce, saltmarsh edge (1 female); beating red spruce foliage (saplings), 

mixed conifer-hardwood (1 female); beating red spruce foliage, mixed conifer (1 female).

Regional Distribution: ME (Procter 1946); NH, MA, CT, NY, ON, QC (Levi 1954a, as E. limbata 

(Walckenaer, 1841)).

Neospintharus trigonum (Hentz, 1850)

Taxonomy: Exline and Levi (1962, as Argyrodes trigonum (Hentz)); Kaston (1981, as Conopistha 

trigona (Hentz)); Paquin and Dupérré (2003, as Argyrodes trigonum (Hentz, 1850)); Agnarrson 

(2004).

Records: M01 (1 male); M06 (1 male); M13 (1 female). N = 3 adults; 2 males, 1 female.

Methods: beating cloth (1 female); search (1 male); sweep net (1 male).

Months: July (1 male); August (1 male, 1 female).

Habitats: beating red spruce foliage, mixed hardwood-conifer (1 female); sweeping vegetation, 

saltmarsh (1 male); in large orb web, timothy grass, old fi eld (1 male).

Regional Distribution: ME (Hentz 1875, as Theridion trigonum); NH, MA, CT, NY, ON (Exline 

and Levi 1962, as Argyrodes trigonum (Hentz)); QC (Bélanger and Hutchinson 1992, as Argyrodes 

trigonum (Hentz)).

Note: This species keys to the genus Argyrodes in Levi (2005c).

Neottiura bimaculata (Linnaeus, 1767)

Taxonomy: Levi (1956); Paquin and Dupérré (2003).

Records: M01 (1 male, 1 female); M07 (1 male); M12 (1 male, 1 female); M14 (1 female). N = 6 

adults; 3 males, 3 females.

Methods: pitfall trap (1 male); search (1 male, 1 female); sweep net (1 male, 2 females).

Months: June (2 males); July (1 male, 2 females); August (1 female).

Habitats: litter, Kalmia-Vaccinium heath (1 male); sweeping grasses and forbs, mixed hardwoodconifer (1 female); sweeping vegetation, freshwater marsh (1 male, 1 female); on bush-honeysuckle 

leaf (1 female); in web, overturned canoe (1 male).
18

Regional Distribution: ME (Jennings, unpubl.); VT (Kochalka 1979); QC (Bélanger and 

Hutchinson 1992). 

Phoroncidia americana (Emerton, 1882)

Taxonomy: Levi (1955b, as Oronota americana (Emerton)); Levi (1964b); Kaston (1981); Paquin 

and Dupérré (2003).

Records: M07 (1 male); M31 (1 male). N = 2 adult males.

Methods: beating cloth (1 male); pitfall trap (1 male).

Months: May (1 male); July (1 male).

Habitats: beating red spruce foliage, mixed white pine-red spruce stand (1 male); litter, KalmiaVaccinium heath (1 male).

Regional Distribution: ME (Jaros-Su et al., unpubl.); MA, CT, NY (Levi 1955b, as Oronota 

americana (Emerton)); NS, ON (Levi 1964b); QC (Bélanger and Hutchinson 1992).

Note: Emerton (1882) fi rst described this species placing it in the genus Ulesanis. Subsequently, 

Levi (1955b) transferred this species to the genus Oronota, and later (Levi 1964b) to the genus 

Phoronicidia.

*Robertus banksi (Kaston, 1946)

Taxonomy: Kaston (1946, as Ctenium banksi n. sp.); Kaston (1981); Paquin and Dupérré (2003).

Records: M12 (5 males, 4 females). N = 9 adults; 5 males, 4 females.

Method: pitfall traps (5 males, 4 females).

Months: June-July (1 male, 1 female); August (1 male, 2 females); August-September (1 male, 1 

female); September (2 males).

Habitat: litter, freshwater marsh edge (5 males, 4 females).

Regional Distribution: ME (this study); NH, VT, MA, NY, ON (Kaston 1946); QC (Bélanger and 

Hutchinson 1992, Paquin and Dupérré 2003).

Rugathodes aurantius (Emerton, 1915)

Taxonomy: Levi (1957b, as Theridion aurantium (Emerton, 1915); Paquin and Dupérré 2003.

Record: M01 (1 female). N = 1 adult female.

Method: pitfall trap (1 female).
19

Month: July (1 female).

Habitat: litter, deciduous-coniferous woodland (1 female).

Regional Distribution: ME (Emerton 1915, as Theridion aurantium Emerton); NY (Crosby and 

Bishop 1928, as Theridion aurantium Emerton); NH, VT, ON, QC, NS, NF (Levi 1957b, as 

Theridion aurantium Emerton).

Rugathodes sexpunctatus (Emerton, 1882)

Taxonomy: Levi (1957b, as Theridion sexpunctatum Emerton, 1882); Kaston (1981, as Theridion 

sexpuncatum Emerton); Paquin and Dupérré (2003).

Records: M01 (2 females). N = 2 adult females.

Method: search (1 female); sweep net (1 female).

Months: May (1 female); June (1 female).

Habitats: sweeping vegetation, freshwater pond edge (1 female); on exterior wall, house (1 female).

Regional Distribution: ME (Bryant 1908, as Theridion sexpunctatus (Emerton)); NY (Crosby and 

Bishop 1928, as Theridion sexpunctatus (Emerton)); NH, VT, MA, ON, QC, NF (Levi 1957b, as 

Theridion sexpunctatum Emerton).

Steatoda americana (Emerton, 1882)

Taxonomy: Levi (1957a, 1962); Kaston (1948, as Asagena americana Emerton, 1882); Kaston 

(1981); Paquin and Dupérré (2003).

Records: M01 (3 males). N = 3 adult males.

Methods: search (3 males).

Months: May (1 male); June (2 males).

Habitats: in hot tub, house deck (1 male); on exterior walls, house (2 males).

Regional Distribution: ME (Bryant 1908, as Asagena americana Emerton); NY (Crosby and Bishop 

1928, as Asagena americana Emerton); NH, VT, RI, CT, ON (Levi 1957a); QC (Bélanger and 

Hutchinson 1992).

Steatoda bipunctata (Linnaeus, 1758)

Taxonomy: Levi (1957a, 1962); Paquin and Dupérré (2003).

Records: M01 (25 males, 39 females); M52 (1 female). N = 65 adults; 25 males, 40 females.
20

Methods: pitfall traps (2 females); search (25 males, 38 females).

Months: February (1 male, 1 female); March (1 male); April (4 females); May (3 males, 12 females); 

June (2 males, 9 females); July (2 males, 5 females); August (6 males, 7 females); September (2 males); 

October (6 males, 1 female); November (2 males, 1 female).

Habitats: litter, Amelanchier-Rubus stand (2 females); in web, base of crabapple tree (1 female); in 

meadowsweet (1 female); in garden (1 female); in webs, outdoor shed (4 females); in webs, house 

basement or cellar (2 males, 11 females); in bathtub, house (9 males, 4 females); in web, bathroom 

wash basin, house (1 male, 1 female); on bathroom wall and ceiling, house (2 females); in webs or 

on interior walls, ceilings, and fl oors, house (6 males, 7 females); in webs or on exterior walls, house 

(1 male, 2 females); in webs or on fl oor, window, storage box, and woodbin, house garage (3 males, 

2 females); on screen door, house (1 female); on patio, house (1 male); in boat (1 male); in water 

bucket, drowned (1 male); in web, interior window frame, barn (1 female).

Regional Distribution: ME (Procter 1938); NH, ON, QC, NB, NS, NF (Levi 1957a).

Note: In some locales along the Atlantic seaboard of New England and in southeastern Canada, this 

invasive species has displaced the native Steatoda borealis (Hentz, 1850) (Nyffeler et al. 1986).

Steatoda borealis (Hentz, 1850)

Taxonomy: Levi (1957a, 1962); Kaston (1981); Paquin and Dupérré (2003).

Records: M01 (1 male, 1 female). N = 2 adults; 1 male, 1 female.

Method: search (1 male, 1 female).

Months: May (1 female); September (1 male).

Habitats: in web, outdoor woodbin (1 male); in rolled garden hose, shed (1 female).

Regional Distribution: ME (Hentz 1875, as Theridion boreale); NY (Crosby and Bishop 1928); 

NH, VT, MA, RI, CT, ON, QC, NS (Levi 1957a). 

Takayus lyricus (Walckenaer, 1842)

Taxonomy: Levi (1957b, as Theridion lyricum Walckenaer); Kaston (1981, as Theridion lyricum 

Walckenaer); Yoshida (2001); Paquin and Dupérré (2003, as Theridion lyricum Walckenaer, 1841).

Record: M01 (1 female). N = 1 adult female.

Method: search (1 female).

Month: July (1 female).

Habitat: on exterior wall, house (1 female).
21

Regional Distribution: ME (Procter 1946, as Theridion kentuckyense Emerton); NY (Crosby and 

Bishop 1928, Theridion kentuckyense Key.); NH, MA, CT, ON (Levi 1957b, as Theridion lyricum

Walckenaer); QC (Bélanger and Hutchinson 1992, as Theridion lyricum Walckenaer, 1841).

Note: This species keys to the genus Theridion in Levi (2005c).

Theridion albidum Banks, 1895

Taxonomy: Levi (1957b, 1963b); Kaston (1981); Paquin and Dupérré (2003).

Records: M01 (1 male, 1 female); M35 (1 male). N = 3 adults; 2 males, 1 female.

Method: sweep net (2 males, 1 female).

Month: July (2 males, 1 female).

Habitats: sweeping vegetation, old fi eld (1 male, 1 female); sweeping marsh vegetation, brackish 

marsh (1 male).

Regional Distribution: ME, MA, CT, NY, ON (Levi 1957b); QC (Bélanger and Hutchinson 1992).

Theridion differens Emerton, 1882

Taxonomy: Levi (1957b, 1963b); Kaston (1981); Paquin and Dupérré (2003).

Records: M01 (4 males, 3 females); M06 (6 males, 10 females); M10 (2 males); M12 (1 female); 

M19 (1 male, 1 female); M28 (1 female); M51 (3 males). N = 32 adults; 16 males, 16 females.

Methods: beating cloth (10 males, 8 females); pitfall traps (1 male, 2 females); search (1 male, 1 

female); sweep net (4 males, 5 females).

Habitats: beating white spruce foliage, saltmarsh edge (4 males, 7 females); beating eastern larch 

foliage (sapling), conifer-mixed hardwood (1 male); beating red spruce foliage, conifer-mixed 

hardwood (1 male); beating red spruce foliage, mixed hardwood-conifer (1 male, 1 female); beating 

eastern larch foliage, mixed conifer opening (2 males); beating red spruce foliage, mixed conifer 

opening (1 male); litter, Amelanchier-Rubus stand (1 male, 2 females); sweeping vegetation, old fi eld 

(2 males); sweeping vegetation (Juncus-Spartina), saltmarsh (1 female); sweeping vegetation, saltmarsh 

edge (2 males, 2 females); sweeping vegetation, freshwater marsh (1 female); sweeping roadside 

vegetation, disturbed land (1 female); in webs, small trees and shrubs (1 female); in woodpile (1 

male).

Regional Distribution: ME (Bryant 1908); NY (Crosby and Bishop 1928); NH, VT, MA, CT, ON, 

QC, NS (Levi 1957b).

Theridion frondeum Hentz, 1850

Taxonomy: Levi (1957b, 1963b); Kaston (1981); Paquin and Dupérré (2003).
22

Records: M01 (3 males, 6 females); M03 (2 females); M04 (1 male, 3 females); M14 (3 males, 4 

females); M23 (4 males); M33 (2 females). N = 28 adults; 11 males, 17 females.

Methods: beating cloth (1 male); search (3 males, 4 females); sweep net (7 males, 11 females); n. d. 

(2 females).

Months: May (2 females); June (3 males, 2 females); July (8 males, 10 females); August (3 females).

Habitats: beating understory red spruce foliage, mixed conifer-hardwood (1 male); sweeping 

Vaccinium and Myrica near woods, old fi eld edge (2 females); sweeping understory vegetation, sprucemixed hardwood (1 female); sweeping roadside grasses, forbs, and shrubs, red spruce forest (1 male); 

sweeping grasses and forbs, mixed hardwood-conifer (1 male, 2 females); sweeping vegetation along 

trail, mixed hardwood-conifer (2 males, 2 females); sweeping Kalmia sp. near freshwater pond, island 

(2 females); sweeping Kalmia, mixed conifer-hardwood, island (1 male); sweeping blueberries, spruce 

(seedlings), mixed conifer opening, island (1 male); sweeping blue-joint grass, seashore-backshore 

(1 male); sweeping marsh grasses and sedges, freshwater marsh (2 females); running across graveled 

lane, mixed hardwood-conifer (2 males, 2 females); in woodpile (2 females); on screen door, house (1 

male); n. d. (2 females).

Regional Distribution: ME (Bryant 1908); NY (Crosby and Bishop 1928); NH, VT, MA, CT, ON, 

QC, NS (Levi 1957b).

Theridion glaucescens Becker, 1879

Taxonomy: Levi (1957b, 1963b); Kaston (1981); Paquin and Dupérré (2003).

Records: M01 (2 females); M06 (2 males); M33 (2 females). N = 6 adults; 2 males, 4 females.

Methods: beating cloth (2 males); search (3 females); sweep net (1 female).

Months: May (2 females); June (2 males, 2 females).

Habitats: beating white spruce, saltmarsh edge (2 males); sweeping grasses and sedges, freshwater 

marsh (1 female); in silk retreat, meadowsweet infl orescence, freshwater marsh edge (1 female); fell 

into hot tub (Jacuzzi), house deck (1 female); on exterior wall, house (1 female).

Regional Distribution: ME (Bryant 1908, as Theridion spirale Emerton); NY (Crosby and Bishop 

1928, as Theridion spirale Emerton); NH, VT, MA, CT, ON, QC, NS, PE, NF (Levi 1957b).

Theridion montanum Emerton, 1882

Taxonomy: Levi (1957b, 1963b); Kaston (1981); Paquin and Dupérré (2003).

Records: M04 (2 females); M48 (1 female). N = 3 adult females.

Methods: beating cloth (1 female); pitfall trap (1 female); sweep net (1 female).
23

Months: August (2 females); September (1 female).

Habitats: beating red spruce foliage, seashore-backshore (1 female); sweeping understory vegetation, 

spruce-mixed hardwood (1 female); litter, coastal red spruce stand (1 female).

Regional Distribution: ME (Emerton 1914); NY (Crosby and Bishop 1928); NH, VT, ON, QC, 

NS, PE, NF (Levi 1957b); NB (Loughton et al. 1963).

Theridion murarium Emerton, 1882

Taxonomy: Levi (1957b, 1963b); Kaston (1981); Paquin and Dupérré (2003).

Records: M01 (5 males, 10 females); M04 (1 female); M06 (2 females); M19 (2 males, 2 females); 

M23 (1 female); M31 (1 female); M51 (4 females). N = 28 adults; 7 males, 21 females.

Methods: beating cloth (4 males, 15 females); pitfall traps (1 male, 2 females); search (2 males, 3 

females); n. d. (1 female).

Months: May (1 male, 2 females); June (3 males, 10 females); July (3 males, 6 females); August (1 

female); September (1 female); October (1 female).

Habitats: beating red spruce foliage, old fi eld edge (2 males, 6 females); beating spruces (1 female); 

beating red spruce foliage, mixed hardwood-conifer (2 females); beating winterberry, roadside, mixed 

hardwood-conifer (1 male); beating balsam fi r foliage, mixed hardwood-conifer (1 male); beating 

red spruce foliage, mixed conifer (2 females); beating red spruce foliage, white pine-red spruce stand 

(1 female); beating eastern larch foliage, mixed conifer (3 females); litter, Amelanchier-Rubus stand 

(1 male); litter, saltmarsh (2 females); freshwater pond edge (1 female); on rock in lawn (1 male); 

running through grass (1 female); in garage, house (1 male); on exterior shingled wall, house (1 

female); n. d., coastal spruce-mixed hardwood (1 female).

Regional Distribution: ME (Bryant 1908); NY (Crosby and Bishop 1928); MA, NH, VT, RI, CT,

ON, QC, NS (Levi 1957b); NB (Loughton et al. 1963).

Theridion pictum (Walckenaer, 1802)

Taxonomy: Levi (1957b, as Theridion ornatum Hahn); Levi 1963b; Kaston (1981, as Theridion 

ornatum Hahn); Paquin and Dupérré (2003).

Records: M02 (2 females); M06 (1 male, 2 females); M09 (1 female); M23 (2 females); M30 (1 

female); M34 (1 female); M50 (1 female). N = 11 adults; 1 male, 10 females.

Methods: beating cloth (2 females); pitfall traps (2 females); search (3 females); sweep net (1 male, 3 

females).

Habitats: beating young balsam fi r foliage, mixed conifer (1 female); beating understory red spruce 

foliage, mixed conifer-hardwood, island (1 female); litter, saltmarsh (2 females); sweeping vegetation, 

saltmarsh edge (1 male); sweeping Kalmia sp., old fi eld, abandoned farm, island (1 female); sweeping 
24

Vaccinium, island (1 female); sweeping grasses, forbs, and shrubs, seashore-backshore (1 female); in 

web, roadside meadowsweet, mixed conifer (1 female); in web, balsam fi r understory, balsam fi r-red 

spruce stand (1 female); in downed burned trees, dry streambed, mixed hardwood-conifer (1 female).

Regional Distribution: ME (Emerton 1882, as Theridion zelotypum Emerton); NY (Crosby and 

Bishop 1928, as Theridion zelotypum Emerton); NH, MA, CT, ON, QC, NS (Levi 1957b, as 

Theridion ornatum Hahn).

Note: During epidemics of the spruce budworm, Choristoneura fumiferana (Clemens), this spider 

captures and feeds on budworm moths entangled in its web (Jennings and Houseweart 1988).

Theridula emertoni Levi, 1954

Taxonomy: Levi (1954b); Kaston (1981); Paquin and Dupérré (2003).

Record: M23 (1 female). N = 1 adult female.

Method: sweep net (1 female).

Month: July (1 female).

Habitat: sweeping Kalmia, mixed conifer opening, island (1 female).

Regional Distribution: ME (Levi 1954b); CT (Kaston 1981); ON, NF (Levi and Randolph 1975); 

QC (Bélanger and Hutchinson 1992).

Thymoites unimaculatus (Emerton, 1882)

Taxonomy: Levi (1957b, as Paidisca unimaculata (Emerton)); Levi (1964a); Kaston (1981, as 

Thymoites unimaculatum (Emerton)); Paquin and Dupérré (2003).

Records: M01 (2 males, 3 females); M05 (1 female); M06 (1 male, 2 females); M12 (1 female); M30 

(1 female); M51 (2 females). N = 13 adults; 3 males, 10 females.

Methods: beating cloth (1 female); pitfall trap (1 female); search (1 female); sweep net (3 males, 7 

females).

Months: May (1 female); June (1 male, 2 females); July (2 males, 4 females); August (3 females).

Habitats: beating eastern larch foliage, mixed conifer (1 female); litter, saltmarsh (1 female); sweeping 

vegetation, old fi elds (1 male, 3 females); sweeping vegetation, freshwater pond edge (1 male); 

sweeping vegetation (Juncus-Spartina), saltmarsh (1 male, 1 female); sweeping marsh grasses and 

sedges, freshwater marsh (1 female); sweeping roadside grasses, forbs, ferns, and rushes, alder-aspen 

edge (1 female); sweeping meadowsweet, mixed conifer opening (1 female); in web, screen house 

interior (1 female).
25

Regional Distribution: ME (Bryant 1908, as Theridion unimaculatum Emerton); NY (Crosby and 

Bishop 1928, as Theridion unimaculatum Em.); MA, CT, RI, NY, ON, QC (Levi 1957b, as Paidisca 

unimaculata Emerton).

Wamba crispulus (Simon, 1895)

Taxonomy: Levi (1957b, as Theridion intervallatum Emerton); Levi (1963b, as Theridion crispulum 

Simon); Kaston (1981, as Theridion crispulum Simon); Wunderlich (1995).

Records: M06 (2 females); M23 (1 female); M31 (1 female). N = 4 adult females.

Methods: beating cloth (2 females); pitfall trap (1 female); n. d. (1 female).

Months: May (1 female); July (3 females).

Habitats: beating white spruce foliage, saltmarsh edge (1 female); beating red spruce foliage (1 

female); litter, saltmarsh (1 female); spruce (Picea sp.), mixed conifer-hardwood, island (1 female).

Regional Distribution: ME (Levi 1957b, as Theridion intervallatum Emerton); NY (Crosby and 

Bishop 1928, as Theridion blandum Hentz); MA (Kaston 1981, as Theridion crispulum Simon); NS 

(Levi and Randolph 1975).

FAMILY LINYPHIIDAE—Subfamily Linyphiinae

The linyphiids or sheet-web spiders are the most diverse family of spiders in Maine. Unlike the 

Theridiidae, the members of this family lack a comb on tarsus IV and usually have thinner legs with 

spines. The chelicerae are generally more robust and lack a boss, but may have a horizontal row of 

striae that forms a stridulating area. The labium is strongly rebordered. Their sheet-like webs usually 

have a platform, bowl, or dome, with an irregular portion above. The resident spider hangs in an 

inverted position below and bites potential prey falling to the bowl or dome.

Some investigators recognize two or more subfamilies of the Linyphiidae; others claim that the 

distinctions between subfamilies are incomplete or lack a consensus among investigators (Buckle 

et al. 2001). Nonetheless, we have chosen to follow Paquin and Dupérré (2003) and recognize two 

prominently represented subfamilies in North America, the Linyphiinae and the Erigoninae. Our 

rationale for doing so is based more on ecological-habitat preferences than on morphological traits. 

In general, most species of Linyphiinae inhabit aerial or aboveground strata, whereas most species of 

Erigoninae are ground or litter inhabitants.

The Linyphiinae are represented by 25 genera and at least 56 species in Maine. Species of Meioneta

(or Agyneta), Lepthyphantes, Oreonetides, Porrhomma, and “unknown” remain to be identifi ed. The 

Linyphiinae fauna at Milbridge consists of 19 genera and 35 species. Unfortunately, identifi cation 

keys are lacking for several species of Linyphiinae in North America. Although a number of genera 

have recently been revised (see Buckle et al. 2001, Draney and Buckle 2005a, and Platnick 2007), 

most have not. Draney and Buckle (2005a) provide an excellent, well-illustrated key to the genera of 

Linyphiinae in North America, as well as family and subfamily character descriptors, notes on natural 

history, and taxonomic history. We include pertinent citations for each of the identifi ed species found 

at Milbridge.
26

*Allomengea dentisetis (Grube, 1861)

Taxonomy: Helsdingen (1974, as Allomengea pinnata (Emerton)); Paquin and Dupérré (2003).

Record: M06 (1 male). N = 1 adult male.

Method: pitfall trap (1 male).

Month: September (1 male).

Habitat: litter, saltmarsh (1 male).

Regional Distribution: ME (this study); NH, MA, NY, NF, NS, NB, ON (Buckle 1988, as 

Allomengea pinnata (Emerton)); QC (Bélanger and Hutchinson 1992).

*Bathyphantes brevis (Emerton, 1911)

Taxonomy: Ivie (1969); Kaston (1981); Paquin and Dupérré (2003).

Record: M01 (1 male). N = 1 adult male.

Method: pitfall trap (1 male).

Month: June-July (1 male).

Habitat: litter, deciduous-coniferous woodland (1 male).

Regional Distribution: ME (this study); VT, MA, CT, NY, ON, NF (Ivie 1969); QC (Bélanger and 

Hutchinson 1992).

Bathyphantes gracilis (Blackwall, 1841)

Taxonomy: Ivie (1969); Paquin and Dupérré (2003).

Records: M12 (2 females). N = 2 adult females.

Method: pitfall traps (2 females).

Months: June (1 female); July (1 female).

Habitat: litter, freshwater marsh edge (2 females).

Regional Distribution: ME (Maloney 2002); NF (Ivie 1969); QC, ON, NS (Buckle et al. 2001).

Bathyphantes pallidus (Banks, 1892)

Taxonomy: Ivie (1969); Kaston (1981); Paquin and Dupérré (2003).
27

Records: M01 (6 males, 3 females); M04 (2 males, 3 females); M12 (32 males, 49 females); M22 

(3 females); M24 (4 males, 5 females); M26 (2 females); M27 (1 female). N = 110 adults; 44 males, 

66 females.

Method: pitfall traps (44 males, 66 females).

Months: June (3 males, 6 females); June-July (4 males, 2 females); July (14 males, 26 females); JulyAugust (9 males, 10 females); August (12 males, 20 females); August-September (1 male, 2 females); 

September (1 male).

Habitats: litter, freshwater marsh edge (32 males, 49 females); litter, red maple sapling stand (4 

males, 5 females); litter, deciduous-coniferous woodland (4 males, 3 females); litter, coastal red spruce 

stand (2 males, 3 females); litter, bigtooth aspen stand (3 females); litter, seashore-backshore (2 

males); litter, saltmarsh edge (2 females); litter, red maple-white (paper) birch stand (1 female).

Regional Distribution: ME, MA, NH, VT, CT, RI, NY, ON, QC, NF, LB (Ivie 1969); NB, NS 

(Buckle et al. 2001).

Centromerus cornupalpis (O. Pickard-Cambridge, 1875)

Taxonomy: Helsdingen (1973); Kaston (1981).

Records: M25 (1 male, 1 female); M27 (1 female). N = 3 adults; 1 male, 2 females.

Method: pitfall traps (1 male, 2 females).

Months: May (1 male); May-June (1 female); July (1 female).

Habitats: litter, white (paper) birch stand (1 male, 1 female); litter, red maple-white (paper) birch 

stand (1 female).

Regional Distribution: ME (Bryant 1908, as Microneta cornupalpis (Cambridge) Emerton); NH, 

VT, MA, CT, NY, NF (Buckle 1988); ON, NS (Buckle et al. 2001).

Centromerus longibulbus (Emerton, 1882)

Taxonomy: Helsdingen (1973); Kaston (1981); Paquin and Dupérré (2003).

Records: M12 (1 male, 1 female). N = 2 adults; 1 male, 1 female.

Method: pitfall traps (1 male, 1 female).

Months: June-July (1 male, 1 female).

Habitat: litter, freshwater marsh edge (1 male, 1 female).

Regional Distribution: ME (Jennings et al. 1988); NH, MA, NY (Helsdingen 1973); QC (Bélanger 

and Hutchinson 1992); NF (Buckle 1988); ON (Buckle et al. 2001).
28

Centromerus persolutus (O. Pickard-Cambridge, 1875)

Taxonomy: Helsdingen (1973); Kaston (1981); Paquin and Dupérré (2003).

Records: M24 (1 male); M25 (1 male); M27 (1 male, 3 females); M53 (7 females). N = 13 adults; 

3 males, 10 females.

Methods: litter condenser-Berlese funnel (7 females); pitfall traps (3 males, 3 females).

Months: May (2 males, 3 females); May-June (1 male); July (7 females).

Habitats: sifted aspen-maple litter, mixed hardwoods (7 females); litter, red maple-white (paper) 

birch stand (1 male, 3 females); litter, red maple sapling stand (1 male); litter, white (paper) birch 

stand (1 male).

Regional Distribution: ME, NH, MA, CT, NY, ON, QC, NF (Helsdingen 1973); NS (Buckle et 

al. 2001).

Centromerus sylvaticus (Blackwall, 1841)

Taxonomy: Helsdingen (1973); Kaston (1981); Paquin and Dupérré (2003).

Records: M01 (15 males, 4 females). N = 19 adults; 15 males, 4 females.

Method: pitfall traps (15 males, 4 females).

Months: October (3 males, 2 females); October-November (12 males, 2 females).

Habitat: litter, edge of old fi eld and alder swamp (15 males, 4 females).

Regional Distribution: ME (Jennings et al., unpubl.); NH, MA, CT, NY, NF, QC, ON (Buckle 

1988).

Diplostyla concolor (Wider, 1834)

Taxonomy: Ivie (1969, as Bathyphantes concolor (Wider)); Kaston (1981, as Bathyphantes concolor 

(Wider)); Paquin and Dupérré (2003, as Bathyphantes concolor (Wider)); Platnick (2007).

Records: M01 (34 males, 5 females); M11 (1 male); M22 (1 male); M26 (1 male). N = 42 adults; 37 

males, 5 females.

Method: pitfall traps (37 males, 5 females).

Months: June (4 males); June-July (5 males, 1 female); July (18 males); July-August (3 males, 

1 female); August (6 males, 2 females); August-September (1 male, 1 female).
29

Habitats: litter, seashore-backshore (25 males, 5 females); litter, deciduous-coniferous woodland near 

seashore (6 males); litter, Amelanchier-Rubus stand near seashore (3 males); litter, bigtooth aspen stand 

(1 male); litter, saltmarsh edge (1 male); ground, gravel pit, mixed conifer-hardwood (1 male).

Regional Distribution: ME, MA, CT, NY, ON, QC, NS, NF (Ivie 1969, as Bathyphantes concolor 

(Wider)).

Note: This species keys to the genus Bathyphantes in Draney and Buckle (2005a); however, see 

Platnick (2007).

Drapetisca alteranda Chamberlin, 1909

Taxonomy: Chamberlin (1909); Kaston (1981); Paquin and Dupérré (2003).

Records: M01 (6 males, 7 females); M04 (1 female). N = 14 adults; 6 males, 8 females.

Methods: brush-tree bole net (2 males, 5 females); search (4 males, 3 females).

Months: August (3 males, 1 female); September (2 males, 6 females); October (1 male, 1 female).

Habitats: brushing bark of white (paper) birch, Betula-Acer stand (2 males, 5 females); on limb of 

apple tree (1 female); on bark of paper birch (1 male); on or under bark of dead tree (1 male), mixed 

hardwood-conifer; on trunk of birch tree, (1 male); spruce-Rugosa, seashore-backshore (1 female); on 

side of house (1 female); on porch post (1 male).

Regional Distribution: ME (Bryant 1908, as Drapetisca socialis (Sundevall) Menge); CT (Kaston 

1981); NY (Crosby and Bishop 1928); QC (Bélanger and Hutchinson 1992); NB (Buckle et al.

2001).

Estrandia grandaeva (Keyserling, 1886)

Taxonomy: Hackman (1954); Kaston (1981); Paquin and Dupérré (2003).

Record: M23 (1 female). N = 1 adult female.

Method: beating cloth (1 female).

Month: July (1 female).

Habitat: beating red spruce foliage, mixed conifer (1 female).

Regional Distribution: ME, NH, MA, NY, NS, NF, LB (Buckle 1988); QC (Bélanger and 

Hutchinson 1992); ON (Buckle et al. 2001).

Frontinella communis (Hentz, 1850)

Taxonomy: Chamberlin and Ivie (1944, as Frontinella pyramitela (Walckenaer,1841)); Kaston 

(1981); Paquin and Dupérré (2003, as Frontinella pyramitela (Walckenaer,1841)).
30

Records: M01 (4 males, 17 females); M04 (1 female); M06 (1 male, 3 females); M07 (1 female); 

M10 (1 male); M14 (2 females); M19 (1 female); M24 (1 female); M51 (1 male, 3 females). N = 36 

adults; 7 males, 29 females.

Methods: beating cloth (1 male, 6 females); pitfall traps (6 females); search (4 males, 13 females); 

sweep net (2 males, 4 females).

Months: May (2 males, 6 females); June (2 males, 13 females); July (3 males, 8 females); August 

(2 females).

Habitats: beating red spruce foliage, seashore-backshore (1 female); beating red spruce foliage, mixed 

hardwood-conifer (1 female); beating red spruce foliage, mixed conifer (1 male, 1 female); beating 

white spruce foliage, saltmarsh edge (1 female); beating Larix laricina foliage, mixed conifer (2 

females); sweeping beach grasses at shore, seashore (1 female); sweeping old fi eld (1 female); sweeping 

grasses, forbs, and shrubs, roadside, red spruce forest (1 female); sweeping saltmarsh vegetation 

(Juncus sp.) (1 male); sweeping low vegetation, mixed conifer-hardwood (1 male); sweeping 

understory vegetation, red maple sapling stand (1 female); litter, Amelanchier-Rubus stand (3 females); 

litter, saltmarsh (2 females); litter, Kalmia-Vaccinium heath (1 female); in webs, dead terminals on 

shrubs, old fi eld-wood edge (2 females); in web among aster blooms (1 female); in web, blackberry 

bush (1 female); in web on alder (1 female); in web on alder along shore, seashore (1 female); in web 

on meadowsweet at shore, seashore (1 female); in web on shrub with female (escaped) (1 male); in 

alders along shore, seashore (1 male); in Prunus (1 male); in web, trailside shrub, mixed hardwoodconifer understory (1 female); along trail, mixed hardwood-conifer understory (1 female); picked 

from shrubs along lane, mixed hardwood-conifer (1 female); in web on common pea vine, vegetable 

garden (1 female); in web on lawn chair (1 female); on cultivated cinquefoil (1 male); on rock, lawn 

(1 female).

Regional Distribution: ME (Bryant 1908, as Linyphia communis Hentz); CT (Kaston 1981); NY 

(Crosby and Bishop 1928, as Linyphia communis Hentz); QC (Bélanger and Hutchinson 1992, as 

Frontinella pyramitela (Walckenaer)).

Notes: Although some recent authors (e.g., Buckle et al. 2001, Paquin and Dupérré 2003, Draney 

and Buckle 2005a) refer to this species as Frontinella pyramitela (Walckenaer, 1841), we follow 

Platnick (2007) who indicates that Frontinella communis (Hentz, 1850) is the valid name for this 

species in North America.

Buckle et al. (2001) noted that F. pyramitela is widespread in Canada and the United States; this 

range no doubt includes all of the New England States and the Maritime Provinces of Canada. 

During outbreaks of the spruce budworm in Maine, this spider captures and feeds on budworm 

moths entangled in its web (Jennings and Houseweart 1988, as F. pyramitela (Walck.)).

Incestophantes calcaratus (Emerton, 1909)

Taxonomy: Zorsch (1937, as Lepthyphantes calcarata (Emerton)); Paquin and Dupérré (2003, as 

Lepthyphantes calcaratus (Emerton, 1909).

Records: M01 (1 male, 2 females); M04 (1 male); M48 (1 female). N = 5 adults; 2 males, 3 females.
31

Methods: beating cloth (1 female); brush-tree bole net (1 female); search (1 female); sweep net 

(2 males).

Months: August (2 males); September (3 females).

Habitats: beating red spruce foliage, seashore-backshore (1 female); brushing bark, white (paper) 

birch, Betula-Acer stand (1 female); sweeping understory vegetation, spruce-mixed hardwood (1 

male); sweeping Vaccinium and small shrubs, old fi eld (1 male); in canoe cover (turned), old fi eld (1 

female).

Regional Distribution: ME, NH, NS (Zorsch 1937, as Lepthyphantes calcaratus (Emerton); QC, NF 

(Buckle 1988, as Lepthyphantes calcaratus (Emerton)); NB, LB (Buckle et al. 2001, as Lepthyphantes 

calcaratus (Emerton 1909)).

Note: This species keys to the genus Lepthyphantes in Draney and Buckle (2005a); however, see 

Platnick (2007).

Incestophantes duplicatus (Emerton, 1913)

Taxonomy: Zorsch (1937, as Lepthyphantes duplicata Emerton); Paquin and Dupérré (2003).

Records: M03 (1 male, 1 female). N = 2 adults; 1 male, 1 female.

Methods: search (1 female); sweep net (1 male).

Month: July (1 male, 1 female).

Habitats: sweeping mixed conifer, freshwater pond edge, island (1 male); in reindeer lichen on ledge, 

mixed conifer-hardwood (1 female).

Regional Distribution: ME (Emerton 1913, as Bathyphantes duplicatus Emerton); QC (Bélanger and 

Hutchinson 1992); NF (Hackman 1954, as Lepthyphantes triramus Chamberlin and Ivie, 1947); NB, 

LB (Buckle et al. 2001, as Lepthyphantes duplicatus Emerton, 1913).

Kaestneria pullata (O. Pickard-Cambridge, 1863)

Taxonomy: Ivie (1969, as Bathyphantes (Coniphantes) pullatus (O. Pickard-Cambridge); Kaston 

(1981, as Bathyphantes pullatus (O. P.-Cambridge)); Paquin and Dupérré (2003).

Records: M12 (1 male, 2 females). N = 3 adults; 1 male, 2 females.

Method: pitfall traps (1 male, 2 females).

Month: July (1 male); July-August (2 females).

Habitats: litter, freshwater marsh edge (1 male, 2 females).
32

Regional Distribution: ME (Dixon, unpubl.); NY (Crosby and Bishop 1928, as Bathyphantes conicus 

Emerton); MA (Kaston 1981, as Bathyphantes pullatus (O. P.-Cambridge); ON, QC (Ivie 1969, as 

Bathyphantes (Coniphantes) pullatus (O. Pickard-Cambridge); NF (Buckle et al. 2001).

Lepthyphantes alpinus (Emerton, 1882)

Taxonomy: (Zorsch 1937, as Lepthyphantes alpina (Emerton)); Paquin and Dupérré (2003).

Records: M12 (1 male, 3 females). N = 4 adults; 1 male, 3 females.

Methods: pitfall traps (1 male, 2 females); sweep net (1 female).

Months: June (1 female); July (1 male); July-August (2 females).

Habitats: litter, freshwater marsh edge (1 male, 2 females); sweeping grasses and other vegetation, 

freshwater marsh (1 female).

Regional Distribution: ME, NH, VT, NY, QC (Zorsch 1937); NF (Buckle 1988); NS (Buckle et al.

2001).

Lepthyphantes leprosus (Ohlert, 1865)

Taxonomy: (Zorsch 1937, as Lepthyphantes leprosa (Ohlert)); (Kaston 1981, as Lepthyphantes leprosa 

(Ohlert)); Paquin and Dupérré (2003).

Records: M01 (1 male, 1 female); M06 (1 male). N = 3 adults; 2 males, 1 female.

Methods: pitfall trap (1 male); search (1 male, 1 female).

Months: May (1 male); June (1 female); October (1 male).

Habitats: litter, saltmarsh (1 male); in web between house door and storm door (1 female); in 

bathroom sink, house (1 male).

Regional Distribution: ME (Procter 1946, as Lepthyphantes leprosa (Ohlert)); NH, RI, NY, NS 

(Zorsch 1937); CT (Kaston 1981, as Lepthyphantes leprosa (Ohlert)); QC (Bélanger and Hutchinson 

1992).

Lepthyphantes turbatrix (O. Pickard-Cambridge, 1877)

Taxonomy: Zorsch (1937, as Lepthyphantes subalpina (Emerton, 1882)); Paquin and Dupérré (2003).

Records: M01 (4 males, 1 female); M04 (1 female); M06 (1 female); M24 (1 female). N = 8 adults; 4 

males, 4 females.

Methods: pitfall traps (1 male, 4 females); search (3 males).

Months: May (1 female); June (2 males); July (1 male, 3 females); August (1 male).
33

Habitats: litter, seashore-backshore (1 male, 1 female); litter, saltmarsh (1 female); litter, coastal red 

spruce stand (1 female); litter, red maple sapling stand (1 female); in house among papers (1 male); 

on bathroom wall, house (1 male); on patio, house (1 male).

Regional Distribution: ME (Jennings et al. 1988); NH, NY, ON, QC, NF (Zorsch 1937); NB 

(Buckle et al. 2001).

Linyphia triangularis (Clerck, 1757)

Taxonomy: Helsdingen (1969); Roberts (1985, 1987).

Records: M01 (5 males, 26 females); M02 (1 female); M04 (1 female); M17 (1 male); M48 (1 

female); M49 (1 male, 1 female); M54 (2 females). N = 39 adults; 7 males, 32 females.

Methods: beating cloth (1 female); search (4 males, 17 females); sweep net (2 males, 13 females); 

shake-sweep net (1 male); n. d. (1 female).

Months: August (4 males, 10 females); September (3 males, 20 females); October (2 females).

Habitats: sweeping vegetation (grasses, forbs, shrubs, small trees), old fi eld (2 males, 13 females); 

beating red spruce foliage, old fi eld edge (1 female); shaking red spruce foliage over sweep net, 

saltmarsh edge (1 male); on red spruce seedling with prey, mixed conifer-hardwood (1 female); 

in webs, small balsam fi r trees, mixed conifer (2 females); in web, small balsam fi r tree, seashorebackshore (1 female); in webs on low understory shrubs, Betula-Acer stand near seashore (2 females); 

in same web on white pine seedling, mixed conifer-hardwood, ledges (1 male, 1 female); in web on 

small trees and shrubs, mixed conifer-hardwood, ledge (1 female); in web on low alder, old fi eld edge 

(1 female); in webs among low wild cherry, old fi eld (1 male, 1 female); in webs on low forbs, old 

fi eld (2 females); in web among mixed low herbs, old fi eld (1 male); in web between two goldenrod 

plants, old fi eld (1 female); in web, old fi eld (1 female); in web, roadside Juncus sp., roadside-mixed 

conifer (1 female); in black crowberry on backshore, seashore-backshore (1 female); among spruceRugosa, seashore-backshore (1 male); in vegetable garden (1 female); n. d. (1 female).

Regional Distribution: ME (Jennings et al. 2002).

Note: This is a Palearctic species recently introduced into coastal regions of Maine (Jennings et al.

2002); thus far, it is not known to occur elsewhere in North America. 

Macrargus multesimus (O. Pickard-Cambridge, 1875)

Taxonomy: Kaston (1981); Paquin and Dupérré (2003).

Records: M01 (3 males, 2 females); M04 (2 males); M25 (3 females). N = 10 adults; 5 males, 5 

females.

Methods: beating cloth (2 males, 1 female); pitfall traps (3 males, 4 females).

Months: May (1 female); May-June (2 males, 2 females); July (1 female); October (3 males, 

1 female).
34

Habitats: litter, white (paper) birch stand (3 females); litter, coastal red spruce stand (2 males); litter, 

edge of old fi eld and alder swamp (1 male); litter, deciduous-coniferous woodland (1 female); beating 

alders and shrubs, old fi eld edge (2 males, 1 female).

Regional Distribution: ME (Procter 1946, as Microneta multesima Cambridge); CT (Kaston 1981); 

NY (Crosby and Bishop 1928, as Microneta multesima Cambridge); MA (Buckle et al. 2001); QC 

(Bélanger and Hutchinson 1992).

Meioneta fabra (Keyserling, 1886)

Taxonomy: Kaston (1981); Saaristo (1973, as Agyneta fabra (Keyserling, 1886)); Saaristo and 

Koponen (1998, as Agyneta fabra (Keyserling, 1886)); Paquin and Dupérré (2003, as Agyneta fabra 

(Keyserling, 1886)); Platnick (2007).

Record: M01 (1 male). N = 1 adult male.

Method: search (1 male).

Month: May (1 male).

Habitat: running through grass near vegetable garden (1 male).

Regional Distribution: ME (Chamberlin 1925, as Bathyphantes wana Chamberlin); CT (Kaston 

1981); NY (Crosby and Bishop 1928, as Microneta fabra Keyserling); QC (Bélanger and Hutchinson 

1992); ON, NB (Buckle et al. 2001, as Agyneta fabra (Keyserling, 1886)).

Note: This species keys to the genus Agyneta in Draney and Buckle (2005a); however, see Platnick 

(2007).

***Meioneta nigripes (Simon, 1884)

Taxonomy: Saaristo and Koponen (1998, as Agyneta nigripes (Simon, 1884)); Paquin and Dupérré 

(2003, as Agyneta nigripes (Simon, 1884)).

Record: M01 (1 female). N = 1 adult female.

Method: pitfall trap (1 female).

Month: July (1 female).

Habitat: litter, mixed deciduous-coniferous woodland (1 female).

Regional Distribution: ME (this study); QC (Paquin and Dupérré 2003, as Agyneta nigripes (Simon, 

1884)).

Note: This species keys to the genus Agyneta in Draney and Buckle (2005a); however, see Platnick 

(2007).
35

Meioneta simplex (Emerton, 1926)

Taxonomy: Kaston (1981); Saaristo and Koponen (1998, as Agyneta simplex (Emerton, 1926)); 

Paquin and Dupérré (2003, as Agyneta simplex (Emerton, 1926)); Platnick (2007).

Records: M01 (1 female); M06 (1 female); M07 (3 males, 1 female); M22 (2 females); M24 (2 

males, 4 females); M25 (2 males, 4 females); M26 (3 males, 1 female); M27 (4 females). N = 28 

adults; 10 males, 18 females.

Method: pitfall traps (10 males, 18 females).

Months: May (1 male); May-June (1 female); June (4 males, 6 females); June-July (1 male, 2 

females); July (4 males, 7 females); July-August (1 female); August (1 female).

Habitats: litter, red maple sapling stand (2 males, 4 females); litter, white (paper) birch stand (2 

males, 4 females); litter, red maple-white birch stand (4 females); litter, bigtooth aspen stand (2 

females); litter, saltmarsh edge (3 males, 1 female); litter, Kalmia-Vaccinium heath (3 males, 1 female); 

litter, Betula-Acer stand (1 female); litter, saltmarsh (1 female).

Regional Distribution: ME (Collins et al. 1996); MA, NS (Emerton 1926, as Lepthyphantes simplex 

n. sp.); QC (Bélanger and Hutchinson 1992); ON (Buckle et al. 2001, as Agyneta simplex (Emerton, 

1926)).

Note: This species keys to the genus Agyneta in Draney and Buckle (2005a); however, see Platnick 

(2007).

Meioneta unimaculata (Banks, 1892)

Taxonomy: Kaston (1981); Paquin and Dupérré (2003, as Agyneta unimaculata (Banks, 1892));

Platnick (2007).

Records: M06 (1 female); M25 (1 male). N = 2 adults; 1 male, 1 female.

Method: pitfall traps (1 male, 1 female).

Months: July (1 male); August (1 female).

Habitats: litter, saltmarsh (1 female); litter, white (paper) birch stand (1 male).

Regional Distribution: ME (Jennings et al., unpubl.); NY (Crosby and Bishop 1928, as 

Bathyphantes unimaculatus Banks); QC (Bélanger and Hutchinson 1992); ON, NS (Buckle et al.

2001, as Agyneta unimaculata (Banks, 1892)).

Note: This species keys to the genus Agyneta in Draney and Buckle (2005a); however, see Platnick 

(2007).
36

Meioneta zygia (Keyserling, 1886)

Taxonomy: Kaston (1981); Platnick (2007).

Records: M12 (2 females); M22 (2 females). N = 4 adult females.

Method: pitfall traps (4 females).

Months: June (1 female); August (3 females).

Habitats: litter, freshwater marsh edge (2 females); litter, bigtooth aspen stand (2 females).

Regional Distribution: ME (Sferra et al., unpubl.); CT (Kaston 1981); NY (Crosby and Bishop 

1928, as Bathyphantes zygios Keyserling); ON (Buckle et al. 2001, as Agyneta zygia (Keyserling, 

1886)).

Note: This species keys to the genus Agyneta in Draney and Buckle (2005a); however, see Platnick 

(2007).

Microlinyphia mandibulata (Emerton, 1882)

Taxonomy: Helsdingen (1970); Kaston (1981); Paquin and Dupérré (2003, as Microlinyphia m. 

mandibulata (Emerton, 1882)).

Records: M01 (1 male, 3 females); M04 (1 female); M06 (1 female); M12 (1 female); M23 (2 

females); M30 (1 female); M33 (1 female). N = 11 adults; 1 male, 10 females.

Methods: search (1 male, 1 female); sweep net (8 females); n. d. (1 female).

Months: May (1 male, 2 females); June (5 females); July (2 females); August (1 female).

Habitats: sweeping old fi eld (1 female); sweeping vegetation, pond edge (1 female); sweeping 

roadside grasses, forbs, and shrubs, red spruce forest (1 female); sweeping saltmarsh vegetation 

(Juncus-Spartina), saltmarsh (1 female); sweeping grasses and shrubs, freshwater marsh (1 female); 

sweeping Vaccinium sp., old fi eld, island (1 female); sweeping roadside vegetation, roadside-alder edge 

(1 female); sweeping understory bordering freshwater marsh, mixed conifer-hardwood (1 female); 

in low plants at edge of lawn (1 female); outside window, house (1 male); spruce, cherry, and grass, 

island (1 female).

Regional Distribution: ME (Emerton 1882); CT (Kaston 1981); NH, NY, NF (Buckle 1988, as 

Microlinyphia mandibulata mandibulata (Emerton 1882); QC (Bélanger and Hutchinson 1992); 

ON, NB (Buckle et al. 2001, as Microlinyphia mandibulata mandibulata (Emerton 1882)).

Microneta viaria (Blackwall, 1841)

Taxonomy: Saaristo (1974); Kaston (1981); Paquin and Dupérré (2003).
37

Records: M01 (1 male, 3 females); M04 (1 male); M24 (6 males); M27 (1 male). N = 12 adults; 9 

males, 3 females.

Methods: pitfall traps (9 males); sifted litter-hand sorted (3 females).

Months: May-June (1 male); June (5 males); June-July (2 males); July (1 male); August (3 females).

Habitats: litter, red maple sapling stand (6 males); litter, red maple-white (paper) birch stand (1 

male); litter, coastal red spruce stand (1 male); litter, seashore-backshore (1 male); sifted litter, 

deciduous-coniferous woodland (3 females).

Regional Distribution: ME (Procter 1946); CT (Kaston 1981); NY (Crosby and Bishop 1928); QC 

(Bélanger and Hutchinson 1992); MA, ON (Buckle et al. 2001).

Neriene clathrata (Sundevall, 1830)

Taxonomy: Helsdingen (1969); Kaston (1981, as Linyphia clathrata (Sundevall)); Paquin and 

Dupérré (2003).

Records: M01 (1 male); M04 (1 female); M06 (1 female); M07 (4 males, 1 female); M14 (1 female); 

M26 (1 female). N = 10 adults; 5 males, 5 females.

Methods: pitfall traps (4 males, 3 females); search (1 male, 1 female); sweep net (1 female).

Months: April (1 male); May (4 males, 2 females); June (2 females); July (1 female).

Habitats: litter, Kalmia-Vaccinium heath (4 males, 1 female); litter, saltmarsh (1 female); litter, 

saltmarsh edge (1 female); sweeping grasses, forbs, and shrubs, roadside, red spruce forest (1 female); 

on ground among leaves and grass, mixed hardwood-conifer (1 female); in garage, house (1 male).

Regional Distribution: ME, NH, MA, CT, NY, NF (Buckle 1988); QC (Bélanger and Hutchinson 

1992); ON, NB, NS (Buckle et al. 2001).

Neriene radiata (Walckenaer, 1842)

Taxonomy: Helsdingen (1969); Kaston (1981, as Prolinyphia marginata (C. L. Koch)); Paquin and 

Dupérré (2003).

Records: M01 (2 females); M04 (2 males, 3 females); M06 (1 female); M07 (2 males, 3 females); 

M14 (1 female); M23 (1 male, 8 females); M34 (1 female); M58 (1 female). N = 25 adults; 5 males, 

20 females.

Methods: pitfall traps (2 males, 4 females); search (9 females); sweep net (3 males, 6 females); n. d. (1 

female).

Months: May (1 female); June (4 males, 7 females); July (1 male, 9 females); August (3 females).
38

Habitats: sweeping grasses, forbs, and shrubs, roadside red spruce forest (2 males, 3 females); 

sweeping grassy area near cabin, mixed conifer opening, island (3 females); sweeping Vaccinium sp., 

island (1 male); litter, Kalmia-Vaccinium heath (2 males, 3 females); litter, saltmarsh (1 female); in 

web on red maple (1 female); in web on red spruce, mixed conifer, island (1 female); in web among 

understory foliage, deciduous-coniferous woodland (1 female); in web, small maple understory, mixed 

hardwood-conifer (1 female); in webs on rock ledges, mixed conifer-hardwood, island (3 females); in 

dome web on ledges, balsam fi r-red spruce-mixed hardwood (1 female); on ground, marsh grasses and 

matted straw, seashore-brackish marsh (1 female); spruce-blueberry-cedar, island (1 female).

Regional Distribution: ME, NH, VT, MA, CT, RI, NY, NS, ON (Buckle 1988); QC (Bélanger and 

Hutchinson 1992); NB (Buckle et al. 2001).

Pityohyphantes costatus (Hentz, 1850)

Taxonomy: Chamberlin and Ivie (1943); Kaston (1981); Paquin and Dupérré (2003).

Records: M01 (7 males, 19 females); M02 (1 female); M04 (2 females); M06 (5 females); M07 (1 

male, 2 females); M10 (1 female); M14 (2 males, 4 females); M23 (1 female); M24 (1 female); M30 

(2 females); M36 (1 male, 1 female); M51 (1 male). N = 51 adults; 12 males, 39 females.

Methods: beating cloth (4 males, 21 females); pitfall traps (1 male, 4 females); search (6 males, 12 

females); sweep net (1 male, 1 female); n. d. (1 female).

Months: April (1 male); May (6 males, 22 females); June (5 males, 13 females); July (3 females); 

August (1 female).

Habitats: beating spruces, old fi eld edge (3 males, 14 females); beating red spruce foliage (1 female); 

beating red spruce foliage, mixed hardwood-conifer understory, forest trail (1 male); beating eastern 

larch foliage, mixed conifer (1 female); beating white spruce foliage, saltmarsh edge (5 females); 

sweeping grasses, forbs, and shrubs, roadside, red spruce forest (1 female); sweeping meadowsweet, 

mixed conifer opening (1 male); litter, coastal red spruce stand (1 female); litter, Kalmia-Vaccinium 

heath (1 male, 2 females); litter, red maple sapling stand (1 female); in webs, shrubs along forest trail, 

mixed hardwood-conifer (1 male, 4 females); in web, balsam fi r understory, balsam fi r-red spruce 

stand (2 females); in same web, dead shrub branchlet, mixed conifer-hardwood (1 male, 1 female); 

under rock on ledge, mixed conifer-hardwood (1 female); in web, cellar window, house (1 female); 

on outside wall, house (2 males); under mulch at edge of lawn (1 male); running in grass (1 female); 

under tarp in screen-house (1 male, 2 females); n. d., island (1 female).

Regional Distribution: ME (Hentz 1875, as Linyphia? costata); CT (Kaston 1981); QC (Bélanger 

and Hutchinson 1992); ON, NS, NF (Buckle et al. 2001).

Pityohyphantes phrygianus (C. L. Koch, 1836)

Taxonomy: Chamberlin and Ivie (1943); Roberts (1985, 1987).

Records: M01 (8 males, 26 females); M07 (1 female); M11 (1 female); M14 (1 male); M15 (8 males, 

14 females); M49 (1 female); M55 (2 males, 2 females). N = 64 adults; 19 males, 45 females.
39

Methods: beating cloth (19 males, 42 females); pitfall traps (2 females); search (1 female).

Months: May (8 males, 15 females); June (11 males, 27 females); July (2 females); August (1 female).

Habitats: beating red spruce foliage, old fi eld edge (3 males, 11 females); beating red spruce foliage 

(5 males, 11 females); beating red spruce foliage, seashore-backshore (3 females); beating red spruce 

foliage, saltmarsh edge (8 males, 14 females); beating red spruce foliage, freshwater streamside (2 

males, 2 females); beating red spruce foliage, mixed hardwood- conifer understory (1 male); beating 

jack pine foliage, mixed conifer-hardwood (1 female); litter, Kalmia-Vaccinium heath (1 female); 

ground, gravel pit, conifer-mixed hardwood (1 female); in shrubs, old fi eld (1 female).

Regional Distribution: ME (Keyserling 1886, as Linyphia phrygiana C. L. Koch); NY (Crosby and 

Bishop 1928); QC (Bélanger and Hutchinson 1992).

*Porrhomma convexum (Westring, 1851)

Taxonomy: Roberts (1985, 1987).

Record: M01 (1 female). N = 1 adult female.

Method: beating cloth (1 female).

Month: June (1 female).

Habitat: beating red spruce foliage, seashore-backshore.

Regional Distribution: ME (this study).

Note: This is a Palaearctic species previously recorded in Alaska, Washington, British Columbia, and 

Greenland (Buckle et al. 2001); it evidently now has been introduced along the northeastern coast of 

Maine.

*Porrhomma terrestre (Emerton, 1882)

Taxonomy: Bishop and Crosby (1938); Chamberlin and Ivie (1947); Kaston (1981); Paquin and 

Dupérré (2003).

Record: M12 (1 male). N = 1 adult male.

Method: pitfall trap (1 male).

Month: August (1 male).

Habitat: litter, freshwater marsh edge (1 male).

Regional Distribution: ME (this study); MA, NY, ON (Buckle 1988); QC (Bélanger and 

Hutchinson 1992).
40

Tapinopa bilineata Banks, 1893

Taxonomy: Chamberlin and Ivie (1944); Kaston (1981); Paquin and Dupérré (2003).

Record: M12 (1 male). N = 1 adult male.

Method: pitfall trap (1 male).

Month: September (1 male).

Habitat: freshwater marsh edge (1 male).

Regional Distribution: ME (Emerton 1909); CT, MA, NY (Kaston 1981); ON, NB, NS (Buckle et 

al. 2001).

Tenuiphantes zebra (Emerton, 1882)

Taxonomy: Zorsch (1937, as Lepthyphantes zebra (Emerton)); Kaston (1981, as Lepthyphantes zebra 

(Emerton)); Saaristo and Tanasevitch (1996); Buckle et al. (2001, as Lepthyphantes zebra (Emerton)); 

Paquin and Dupérré (2003).

Records: M01 (1 male); M12 (4 females); M24 (3 females); M27 (1 female). N = 9 adults; 1 male, 

8 females.

Methods: pitfall traps (8 females); search (1 male).

Months: June (3 females); June-July (1 female); July (1 female); September (1 male, 3 females).

Habitats: litter, freshwater marsh edge (4 females); litter, red maple sapling stand (3 females); litter, 

red maple-white (paper) birch stand (1 female); in overturned canoe, old fi eld (1 male).

Regional Distribution: ME (Bryant 1908, as Lepthyphantes zebra Emerton); MA, CT, VT, NY, ON, 

QC (Buckle 1988, as Lepthyphantes zebra (Emerton)); NB, NS (Buckle et al. 2001, as Lepthyphantes 

zebra (Emerton)).
41

FAMILY LINYPHIIDAE—Subfamily Erigoninae

The subfamily Erigoninae was formerly given full family status; fi rst as the Micryphantidae, then 

later as the Erigonidae. Modern investigators have incorporated these earlier families into the 

Linyphiidae, with their member genera and species assigned to the subfamily Erigoninae or other 

subfamilies. However, controversy remains about the distinction and characterization of some 

linyphiid subfamilies (see Buckle et al. 2001). We follow Paquin and Dupérré (2003) in recognizing 

both the Linyphiinae and Erigoninae; most North American linyphiids belong to either of these two 

subfamilies.

Members of the Erigoninae are generally smaller than their counterparts assigned to the Linyphiinae. 

In general, erigonids are usually ground dwellers that live in litter on the forest fl oor, or in 

saltmarshes, freshwater marshes, seashores, and other habitats. A few species are arboreal.

Unfortunately, identifi cation keys are generally lacking for most genera and species of Erigoninae; 

however, recently Draney and Buckle (2005b) published a generic key for identifying male erigonids. 

Although limited to males, this generic key is a gigantic leap forward in spider identifi cation 

technology. The erigonid fauna in Maine comprises some 42 genera and 115 species; thus far, 

29 genera (1 unknown) and 54 species (2 unknown) have been found at Milbridge. We cite only 

pertinent literature for each of the species found in Milbridge; see Buckle et al. (2001) for a more 

detailed review of literature and Platnick (2007) for recent name changes.

Baryphyma trifrons (O. Pickard-Cambridge, 1863)

Taxonomy: Crosby and Bishop (1933, as Minyriolus aquatilis Crosby & Bishop)); Buckle et al.

(2001, as Baryphyma trifrons affi ne (Schenkel, 1930)); Paquin and Dupérré (2003, as Baryphyma 

trifrons affi ne (Schenkel, 1930).

Records: M01 (1 female); M05 (4 males, 8 females); M07 (3 females); M12 (12 females); M23 (1 

female); M33 (5 females). N = 34 adults; 4 males, 30 females.

Methods: pitfall traps (4 females); search (1 female); sweep net (4 males, 25 females).

Months: May (4 males, 8 females); June (14 females); July (7 females); September (1 female).

Habitats: litter, Kalmia-Vaccinium heath (3 females); litter, freshwater marsh edge (1 female); 

sweeping vegetation, old fi eld (1 female); sweeping grasses and forbs in open fi eld, old fi eld (4 males, 

8 females); sweeping grasses and sedges, freshwater marshes (5 females); sweeping grasses and other 

vegetation, freshwater marsh (9 females); sweeping marsh grasses, freshwater marsh (2 females); in 

grass along beach, island, seashore (1 female).

Regional Distribution: ME (Mairs and Jennings, unpubl.); NY (Carroll and Roth 1988, as 

Baryphyma trifrons affi ne (Schenkel, 1930)); QC (Bélanger and Hutchinson 1992).

Note: According to Buckle et al. (2001), the subspecies trifrons affi ne apparently has not been 

formally reduced to full synonymy with trifrons. Some recent authors (e.g., Aitchison-Benell and 
42

Dondale 1992, Bélanger and Hutchinson 1992, Eskov 1994) have used trifrons rather than trifrons 

affi ne; however, see Paquin and Dupérré (2003).

Carorita limnaea (Crosby & Bishop, 1927)

Taxonomy: Crosby and Bishop (1927); Duffey and Merrett (1964); Paquin and Dupérré (2003).

Record: M12 (1 male). N = 1 adult male.

Method: pitfall trap (1 male).

Month: June-July (1 male).

Habitat: freshwater marsh edge (1 male).

Regional Distribution: ME (Jennings et al. 1988, as Carorita limnaeus (Crosby & Bishop)); NY, 

ON, QC (Buckle et al. 2001).

Ceraticelus bulbosus (Emerton, 1882)

Taxonomy: Crosby and Bishop (1925); Kaston (1981); Paquin and Dupérré (2003).

Records: M05 (2 males). N = 2 adult males.

Method: sweep net (2 males).

Month: May (2 males).

Habitat: sweeping grasses and forbs, old fi eld (2 males).

Regional Distribution: ME (Hilburn and Jennings 1988); CT, NY, ON, QC, NF (Buckle et al.

2001).

Ceraticelus emertoni (O. Pickard-Cambridge, 1874)

Taxonomy: Crosby and Bishop (1925); Kaston (1981); Paquin and Dupérré (2003).

Record: M01 (1 male). N = 1 adult male.

Method: sweep net (1 male).

Month: September (1 male).

Habitat: sweeping grasses, forbs, shrubs, and small trees, old fi eld (1 male).

Regional Distribution: ME (Bryant 1908, as Ceratinella emertoni (Cambridge) Emerton); NH, MA, 

CT, NY, ON, QC, NS (Buckle et al. 2001).
43

Ceraticelus fi ssiceps (O. Pickard-Cambridge, 1874)

Taxonomy: Crosby and Bishop (1925); Kaston (1981); Paquin and Dupérré (2003).

Records: M01 (5 females); M07 (1 female); M23 (1 female); M27 (1 female). N = 8 adult females.

Methods: beating cloth (1 female); pitfall traps (2 females); search (1 female); sweep net (4 females).

Months: May (1 female); June (1 female); July (2 females); August (4 females).

Habitats: beating red spruce foliage (1 female); litter, Kalmia-Vaccinium heath (1 female); litter, 

red maple-white (paper) birch stand (1 female); sweeping Vaccinium and small shrubs (2 females); 

sweeping aster and raspberry, old fi eld (1 female); sweeping grassy area around cabin, mixed conifer 

opening, island (1 female); under white (paper) birch bark, Betula-Acer stand (1 female).

Regional Distribution: ME (Bryant 1908, as Ceratinella fi ssiceps (Cambridge) Emerton); NH, MA, 

CT, NY, ON, QC, NB, NS, NF (Buckle et al. 2001).

Ceraticelus laetabilis (O. Pickard-Cambridge, 1874)

Taxonomy: Crosby and Bishop (1925); Kaston (1981); Paquin and Dupérré (2003).

Records: M24 (4 males, 1 female); M25 (7 males); M26 (1 male); M27 (4 males). N = 17 adults; 16 

males, 1 female.

Method: pitfall traps (16 males, 1 female).

Months: May (1 female); June (15 males); July (1 male).

Habitats: litter, red maple sapling stand (4 males, 1 female); litter, white (paper) birch stand (7 

males); litter, saltmarsh edge (1 male); litter, red maple-white birch stand (4 females).

Regional Distribution: ME (Jennings et al. 1988); NH, VT, MA, CT, NY, ON, QC, NB, NS, NF 

(Buckle et al. 2001).

Ceraticelus laticeps (Emerton, 1894)

Taxonomy: Crosby and Bishop (1925); Kaston (1981).

Record: M07 (1 male). N = 1 adult male.

Method: unsifted litter-Berlese funnel (1 male).

Month: August (1 male).

Habitat: litter, Kalmia-Vaccinium heath (1 male).

Regional Distribution: ME (Jennings et al., unpubl.); NY, ON (Buckle et al. 2001).
44

Ceraticelus minutus (Emerton, 1882)

Taxonomy: Crosby and Bishop (1925); Kaston (1981); Paquin and Dupérré (2003).

Records: M24 (1 female); M53 (2 females). N = 3 adult females.

Methods: litter condenser-Berlese funnel (2 females); pitfall trap (1 female).

Months: May-June (1 female); July (2 females).

Habitats: aspen-maple litter, mixed hardwoods (2 females); litter, red maple sapling stand (1 female).

Regional Distribution: ME (Jennings et al. 1988); NH, MA, CT, NY, ON, QC (Buckle et al.

2001).

Ceratinella brunnea Emerton, 1882

Taxonomy: Crosby and Bishop (1925); Kaston (1981); Paquin and Dupérré (2003).

Records: M04 (5 females); M24 (1 female); M26 (1 female). N = 7 adult females.

Method: pitfall traps (7 females).

Months: June (4 females); July (2 females); July-August (1 female).

Habitats: litter, coastal red spruce stand (5 females); litter, red maple sapling stand (1 female); litter, 

saltmarsh edge (1 female).

Regional Distribution: ME (Procter 1946); NH, CT, NY, ON, QC, NB, NF, LB (Buckle et al.

2001).

Ceratinella parvula (Fox, 1891)

Taxonomy: Crosby and Bishop (1925); Kaston (1981); Paquin and Dupérré (2003).

Record: M35 (1 female). N = 1 adult female.

Method: litter condenser-Berlese funnel (1 female).

Month: June (1 female).

Habitat: marsh litter (grasses, rushes), brackish marsh (1 female).

Regional Distribution: ME (Mairs and Jennings, unpubl.); MA, NY, ON, QC (Buckle et al. 2001).

*Ceratinops annulipes (Banks, 1892)

Taxonomy: Crosby and Bishop (1933); Kaston (1981); Paquin and Dupérré (2003).
45

Records: M26 (1 female); M56 (17 males, 19 females). N = 37 adults; 17 males, 20 females.

Methods: litter condenser-Berlese funnel (9 males, 7 females); litter condenser-hand sorted (8 males, 

12 females); pitfall trap (1 female).

Months: July (1 female); September (17 males, 19 females).

Habitats: litter, saltmarsh edge (1 female); litter (grasses, forbs, tidal debris), salt meadow (17 males, 

19 females).

Regional Distribution: ME (this study); NY, ON, QC (Buckle et al. 2001).

Ceratinopsis nigriceps Emerton, 1882

Taxonomy: Bishop and Crosby (1930); Kaston (1981); Paquin and Dupérré (2003).

Records: M04 (1 female); M06 (1 male); M07 (1 female); M23 (1 female). N = 4 adults; 1 male, 

3 females.

Methods: beating cloth (1 male); pitfall trap (1 female); sweep net (2 females).

Months: June (2 females); July (1 male, 1 female).

Habitats: beating white spruce foliage, saltmarsh edge (1 male); litter, Kalmia-Vaccinium heath (1 

female); sweeping Vaccinium sp., mixed conifer-hardwood, island (1 female); sweeping understory 

grasses, forbs, and shrubs, red spruce forest (1 female).

Regional Distribution: ME (Murray and Jennings, unpubl.); NH, MA, CT, NY, ON, QC, NB, NS 

(Buckle et al. 2001).

Collinsia plumosa (Emerton, 1882)

Taxonomy: Crosby and Bishop (1928, as Catabrithorax plumosus (Emerton) (male only); Kaston 

(1981, as Collinsia oxypaederotipus (Crosby)); Paquin and Dupérré (2003, as Halorates plumosus 

(Emerton, 1882)); Buckle et al. (2001, as Halorates plumosus (Emerton, 1882)); Platnick (2007).

Records: M01 (1 female); M06 (1 male); M26 (1 male); M35 (1 female). N = 4 adults; 2 males, 

2 females.

Methods: pitfall traps (2 males); search (1 female); sweep net (1 female).

Months: May (1 male, 1 female); July (1 male, 1 female).

Habitats: litter, saltmarsh (1 male); litter, saltmarsh edge (1 male); sweeping vegetation, brackish 

marsh (1 female); on large rock, seashore (1 female).
46

Regional Distribution: ME (Hilburn and Jennings 1988, as Halorates plumosus (Emerton, 1882)); 

NH, MA, CT, NY, ON, QC, NS, NB (Buckle et al. 2001, as Halorates plumosus (Emerton, 1882)).

Note: Some authors place this species in the genus Halorates; others place it in the genus Collinsia. 

See the discussion in Buckle et al. (2001) and subsequent listing by Platnick (2007).

Diplocephalus cristatus (Blackwall, 1833)

Taxonomy: Bishop and Crosby (1935); Kaston (1981); Paquin and Dupérré (2003).

Records: M01 (2 males, 11 females). N = 13 adults; 2 males, 11 females.

Method: pitfall traps (2 males, 11 females).

Months: June (1 male, 7 females); June-July (3 females); July (1 male, 1 female).

Habitat: litter, seashore-backshore (2 males, 11 females).

Regional Distribution: ME (Procter 1933); MA, ON, QC, NF (Buckle et al. 2001).

Diplocephalus subrostratus (O. Pickard-Cambridge, 1873)

Taxonomy: Crosby and Bishop (1933); Paquin and Dupérré (2003).

Records: M01 (1 female); M04 (2 males, 1 female). N = 4 adults; 2 males, 2 females.

Methods: pitfall traps (2 males, 1 female); search (1 female).

Months: June (2 males, 1 female); July (1 female).

Habitats: litter, coastal red spruce stand (2 males, 1 female); on deck, house (1 female).

Regional Distribution: ME (Procter 1946, as Chocorua cuneata Emerton); NH, NY, ON, QC, NS, 

NF (Buckle et al. 2001).

Dismodicus decemoculatus (Emerton, 1882)

Taxonomy: Crosby and Bishop (1933); Paquin and Dupérré (2003).

Record: M23 (1 female).

Method: sweep net (1 female).

Month: July (1 female).

Habitat: sweeping Kalmia sp. near freshwater pond, mixed conifer-hardwood, island (1 female).

Regional Distribution: ME (Jennings and Hilburn 1988, as Dismodicus bifrons decemoculatus 

(Emerton, 1882)); NH, NY, ON, QC, NB, NS, NF (Buckle et al. 2001).
47

Eperigone entomologica (Emerton, 1911)

Taxonomy: Crosby and Bishop (1928); Kaston (1981); Millidge (1987); Paquin and Dupérré (2003).

Record: M54 (1 female). N = 1 adult female.

Method: litter condenser-Berlese funnel (1 female).

Month: September (1 female).

Habitat: sifted forest-fl oor litter, red spruce stand (1 female).

Regional Distribution: ME, MA, CT, NY, ON, QC (Millidge 1987, Buckle et al. 2001).

*Eperigone undulata (Emerton, 1914)

Taxonomy: Crosby and Bishop (1928, as Eperigone contorta var. undulatum); Millidge (1987); Paquin 

and Dupérré (2003).

Record: M53 (1 male). N = 1 adult male.

Method: litter condenser-Berlese funnel (1 male).

Month: July (1 male).

Habitat: sifted aspen-maple litter, mixed hardwoods (1 male).

Regional Distribution: ME (this study); NY, ON, QC, NB, NS (Millidge 1987, Buckle et al. 2001).

Erigone aletris Crosby & Bishop, 1928

Taxonomy: Crosby and Bishop (1928); Kaston (1981); Paquin and Dupérré (2003).

Records: M01 (2 males, 5 females). N = 7 adults; 2 males, 5 females.

Methods: litter condenser-Berlese funnel (4 females); pitfall traps (2 males, 1 female).

Months: June (1 male, 4 females); July (1 male); August (1 female).

Habitats: litter, seashore-backshore (2 males, 1 female); sifted rockweed (Fucus vesiculosis) debris, 

upper shore, seashore-backshore (4 females).

Regional Distribution: ME (Crosby and Bishop 1928); MA, NY, QC, NF (Buckle et al. 2001).

Erigone atra Blackwall, 1833

Taxonomy: Crosby and Bishop (1928); Kaston (1981); Paquin and Dupérré (2003).

Records: M01 (1 male); M06 (1 male); M12 (1 male). N = 3 adult males.
48

Methods: beating cloth (2 males); sweep net (1 male).

Months: May (1 male); June (2 males).

Habitats: beating spruce foliage (1 male); beating white spruce foliage, saltmarsh edge (1 male); 

sweeping marsh grasses and shrubs (Spiraea-Kalmia), freshwater marsh (1 male).

Regional Distribution: ME (Bryant 1908, as Erigone longipalpis (Sundevall) Menge); NH, MA, CT, 

NY, ON, QC, NB, NS, NF (Buckle et al. 2001).

*Erigone blaesa Crosby & Bishop, 1928

Taxonomy: Crosby and Bishop (1928); Kaston (1981); Paquin and Dupérré (2003).

Records: M01 (5 males, 2 females); M07 (1 male). N = 8 adults; 6 males, 2 females.

Methods: pitfall traps (3 males, 1 female); search (3 males, 1 female).

Months: June (3 males, 1 female); July (1 male); October (1 male); November (1 male, 1 female).

Habitats: litter, seashore-backshore (1 male, 1 female); litter, mixed deciduous-coniferous woodland 

(1 male); litter, Kalmia-Vaccinium heath (1 male); on rocks at shore, seashore (1 male); running 

swiftly among wet rocks and sand, seashore (1 male); under board, cobble beach, seashore (1 male, 

1 female).

Regional Distribution: ME (this study); MA, CT, RI, NY, ON, QC, NF, LB (Buckle et al. 2001).

***Erigone dentipalpis (Wider, 1834)

Taxonomy: Locket and Millidge (1953); Roberts (1987).

Records: M07 (1 female); M33 (1 female); M50 (1 male). N = 3 adults; 1 male, 2 females.

Methods: pitfall trap (1 female); sweep net (1 male, 1 female).

Months: June (2 females); August (1 male).

Habitats: litter, Kalmia-Vaccinium heath (1 female); sweeping grasses and sedges, freshwater marsh 

(1 female); sweeping grasses, forbs, and shrubs, seashore-backshore (1 male).

Regional Distribution: ME (this study); NF (Buckle et al. 2001).

Note: Buckle et al. (2001) noted that this Palearctic species was possibly introduced into 

Newfoundland. Our collections of Erigone dentipalpis in Milbridge provide the fi rst records of this 

species in New England and in the continental United States.
49

Erigone ephala Crosby & Bishop, 1928

Taxonomy: Crosby and Bishop (1928); Kaston (1981).

Record: M23 (1 female). N = 1 adult female.

Method: sweep net (1 female).

Month: July (1 female).

Habitat: sweeping blue-joint grass, seashore-backshore, island (1 female).

Regional Distribution: ME (Crosby and Bishop 1928); MA, NY, NF (Buckle et al. 2001).

*Erigone zographica Crosby & Bishop, 1928

Taxonomy: Crosby and Bishop (1928); Paquin and Dupérré (2003).

Records: M01 (2 males). N = 2 adult males.

Methods: pitfall traps (1 male); search (1 male).

Months: June (1 male); July (1 male).

Habitats: litter, seashore-backshore (1 male); in high saltgrass (Spartina sp.) at shore, seashorebackshore (1 male).

Regional Distribution: ME (this study); NY, QC (Buckle et al. 2001).

*Floricomus rostratus (Emerton, 1882)

Taxonomy: Bishop and Crosby (1935); Kaston (1981); Paquin and Dupérré (2003).

Records: M26 (2 males). N = 2 adult males.

Method: pitfall traps (2 males).

Months: May-June (1 male); July (1 male).

Habitat: litter, saltmarsh edge (2 males).

Regional Distribution: ME (this study); MA, CT, ON, QC (Buckle et al. 2001).

Gonatium crassipalpum Bryant, 1933

Taxonomy: Bishop and Crosby (1938, as Gonatium rubens (Blackwall)); Kaston (1981, as Gonatium 

rubens (Blackwall)); Millidge (1981a); Paquin and Dupérré (2003).

Records: M01 (1 male, 1 female). N = 2 adults; 1 male, 1 female.
50

Method: sweep net (1 male, 1 female).

Month: September (1 male, 1 female).

Habitats: sweeping grasses, forbs, shrubs, and small trees, old fi eld (1 male); sweeping vegetation, old 

fi eld (1 female).

Regional Distribution: ME (Bishop and Crosby 1935, as Gonatium rubens (Blackwall)); NH, VT, 

MA, CT, RI, NY, ON, QC, NB, NS, LB (Buckle et al. 2001).

Note: According to Buckle et al. (2001), early North American records of Gonatium rubens are 

misidentifi cations of G. crassipalpum. In his revision of Gonatium, Millidge (1981a) established that 

G. crassipalpum was distinct from the Palearctic G. rubens.

Grammonota angusta Dondale, 1959

Taxonomy: Dondale (1959); Paquin and Dupérré (2003).

Records: M01 (18 males, 40 females); M03 (1 female); M04 (6 females); M06 (9 females); M07 (1 

male); M08 (1 male); M10 (3 males, 5 females); M11 (8 females); M12 (1 female); M13 (4 females); 

M14 (5 males, 2 females); M15 (6 males, 6 females); M19 (1 female); M23 (1 male, 1 female); M31 

(5 females); M47 (1 male); M48 (1 female); M49 (2 males, 5 females); M51 (1 female); M55 (3 

females). N = 137 adults; 38 males, 99 females.

Methods: beating cloth (33 males, 86 females); pitfall traps (1 male, 9 females); search (1 male); 

sifted litter-hand sorted (1 female); sweep net (3 males, 2 females); n. d. (1 female).

Months: May (20 males, 32 females); June (15 males, 24 females); July (3 males, 27 females); August 

(10 females); September (1 female); October (5 females).

Habitats: beating red spruce foliage, old fi eld edge (16 males, 24 females); beating red spruce foliage, 

coastal spruce-mixed hardwood (5 females); beating red spruce foliage (saplings), conifer-mixed 

hardwood (3 males, 3 females); beating red spruce foliage, mixed hardwood-conifer (4 females); 

beating red spruce foliage, mixed conifer-hardwood understory (5 males, 2 females); beating red 

spruce foliage, saltmarsh edge (6 males, 6 females); beating red spruce foliage, white pine-red spruce 

stand (5 females); beating red spruce foliage, seashore-upper shore (2 males, 5 females); beating red 

spruce foliage, mixed conifer opening (1 female); beating red spruce foliage, freshwater streamside, 

riparian (3 females); beating red spruce foliage (13 females); beating white spruce foliage, saltmarsh 

edge (9 females); beating spruces (3 females); beating jack pine foliage (1 male); beating eastern larch 

foliage (saplings), mixed conifer-hardwood (2 females); beating balsam fi r foliage, mixed hardwoodconifer (1 female); sweeping grasses, forbs, and sweetpea, seashore-upper beach (1 male); sweeping 

grasses, freshwater marsh edge (1 female); sweeping mixed conifer, freshwater pond edge, island (1 

female); sweeping Vaccinium sp., mixed conifer-hardwood, island (1 male); sweeping vegetation, 

freshwater streamside, riparian (1 male); sifting forest-fl oor litter under maples (Acer sp.), mixed 

hardwood-conifer (1 female); litter, coastal red spruce stand (1 female); litter, Kalmia-Vaccinium heath 

(1 male); ground, gravel pit, conifer-mixed hardwood opening (8 females); cherry, mixed coniferhardwood, island (1 female); on wall, house (1 male).
51

Regional Distribution: ME (Dondale 1959); NY, ON, QC, NB, NS, PE, NF, LB (Buckle et al.

2001).

Grammonota gigas (Banks, 1896)

Taxonomy: Bishop and Crosby (1933); Kaston (1981); Paquin and Dupérré (2003).

Records: M01 (1 female); M06 (10 males, 3 females); M12 (5 females); M26 (9 males, 1 female); 

M35 (3 males, 4 females); M56 (11 females). N = 47 adults; 22 males, 25 females.

Methods: litter condenser-Berlese funnel (3 males, 10 females); litter condenser-hand sorted 

(5 females); pitfall traps (19 males, 8 females); search (1 female); sweep net (1 female).

Months: June (5 males, 5 females); July (15 males, 3 females); July-August (2 males, 2 females); 

August (4 females); September (11 females).

Habitats: litter, saltmarsh (10 males, 3 females); litter, saltmarsh edge (9 males); litter, freshwater 

marsh edge (5 females); sifted marsh litter (grasses, rushes), brackish marsh (3 males, 4 females); sifted 

litter (grasses, forbs, tidal debris), salt meadow (11 females); sweeping grasses, saltmarsh (1 female); 

on cobble beach under stone, seashore (1 female).

Regional Distribution: ME (Hilburn and Jennings 1988); NH, MA, NY, ON, QC, NF (Buckle et 

al. 2001).

***Grammonota maritima Emerton, 1925

Taxonomy: Bishop and Crosby (1933); Paquin and Dupérré (2003).

Records: M56 (2 males, 3 females). N = 5 adults; 2 males, 3 females.

Method: litter condenser-hand sorted (2 males, 3 females).

Month: September (2 males, 3 females).

Habitat: sifted litter (grasses, forbs, tidal debris), salt meadow (2 males, 3 females).

Regional Distribution: ME (this study); QC, NS, NF (Buckle et al. 2001).

Grammonota trivittata Banks, 1895

Taxonomy: Bishop and Crosby (1933); Dondale (1959); Kaston (1981).

Records: M05 (2 females); M17 (1 female). N = 3 adult females.

Methods: sifted litter-hand sorted (2 females); sweep net (1 female).

Month: August (3 females).
52

Habitats: sifted dried rockweed, old fi eld-brackish marsh edge (2 females); sweeping vegetation, 

saltmarsh (1 female).

Regional Distribution: ME (Emerton 1914); MA, CT, RI, NY (Buckle et al. 2001).

Hypselistes fl orens (O. Pickard-Cambridge, 1875)

Taxonomy: Crosby and Bishop (1933); Kaston (1981); Paquin and Dupérré (2003).

Records: M01 (11 females); M04 (4 females); M05 (1 female); M06 (26 females); M07 (12 females); 

M08 (5 females); M12 (4 females); M14 (13 females); M17 (1 female); M19 (2 females); M23 (2 

females); M24 (4 females); M26 (2 females); M28 (1 female); M33 (4 females); M34 (2 females); 

M35 (6 females); M37 (3 females); M47 (2 females); M51 (1 female); M55 (3 females); M57 (2 

females); M58 (3 females). N = 114 adult females.

Methods: beating cloth (5 females); pitfall traps (23 females); search (8 females); sweep net (78 

females).

Months: May (11 females); June (53 females); July (35 females) August (15 females).

Habitats: beating white spruce foliage, saltmarsh edge (1 female); beating winterberry, roadside, 

mixed hardwood-conifer (1 female); beating wild-raisin, brackish marsh edge (1 female); beating 

meadowsweet near streamside, riparian (2 females); sweeping Vaccinium and Myrica near woods, 

old fi eld edge (2 females); sweeping low shrubs, understory Betula-Acer woods (2 females); sweeping 

vegetation, freshwater pond edge (3 females); sweeping stand of meadowsweet (1 female); sweeping 

vegetation, old fi elds (3 females); sweeping vegetation, low wet area, coastal red spruce-mixed 

hardwood (1 female); sweeping vegetation (Juncus-Spartina), saltmarsh (3 females); sweeping 

vegetation, saltmarsh (10 females); sweeping vegetation, saltmarsh edge (1 female); sweeping grasses, 

forbs, and seaside-pea, seashore-upper beach (3 females); sweeping grasses and shrubs on shore, 

seashore (2 females); sweeping grasses, saltmarsh (1 female); sweeping vegetation, saltmarsh edge 

(2 females); sweeping grasses and sedges, freshwater marsh (2 females); sweeping grasses and other 

vegetation, freshwater marsh (2 females); sweeping grasses and forbs, forest trail, mixed hardwoodconifer (9 females); sweeping vegetation along trail, mixed hardwood-conifer (4 females); sweeping 

mixed hardwoods and conifers, roadside mixed hardwood-conifer (1 female); sweeping blue-joint 

grass, seashore-backshore, island (1 female); sweeping understory vegetation, white birch stand (2 

females); sweeping understory vegetation, red maple sapling stand (2 females); sweeping disturbed 

land, roadside (1 female); sweeping grasses and sedges, freshwater marsh (3 females); sweeping 

marsh grasses, freshwater marsh (1 female); sweeping vegetation, dry streambed, mixed hardwoodconifer (1 female); sweeping grasses, sedges, and forbs, dry streambed, mixed hardwood-conifer 

(1 female); sweeping vegetation, brackish marsh (4 females); sweeping marsh grasses and rushes, 

brackish marsh (1 female); sweeping vegetation, mown fi eld edge (3 females); sweeping vegetation, 

freshwater streamside, riparian (2 females); sweeping rhodora and Spiraea, mixed conifer opening 

(1 female); sweeping mixed vegetation, school playground edge, disturbed area (1 female); sweeping 

rushes, brackish marsh (2 females); litter, saltmarsh (11 females); litter, Kalmia-Vaccinium heath (12 

females); on rocks at shore, seashore (1 female); on cobblestone beach and ledge, seashore (1 female); 

on reindeer lichen, ledge near seashore (1 female); on ledge under fl aked stone, mixed conifer, island 
53

(1 female); on ground near brackish marsh, seashore-brackish marsh (2 females); on ground, grass 

and matted straw, seashore-brackish marsh (1 female); in window, house (1 female).

Regional Distribution: ME (Procter 1946); NH, MA, CT, NY, QC, ON, NB, NS (Buckle et al.

2001).

*Islandiana fl aveola (Banks, 1892)

Taxonomy: Bishop and Crosby (1936); Ivie (1965); Kaston (1981); Paquin and Dupérré (2003).

Records: M01 (1 male); M06 (3 males). N = 4 adult males.

Method: pitfall traps (4 males).

Months: May (1 male); June (1 male); July (1 male); August (1 male).

Habitats: litter, seashore-backshore (1 male); litter, saltmarsh (3 males).

Regional Distribution: ME (this study); MA, NY, ON, QC (Buckle et al. 2001).

Metopobactrus prominulus (O. Pickard-Cambridge, 1872)

Taxonomy: Bishop and Crosby (1935, as Maso alticeps Emerton); Kaston (1981, as Maso alticeps

(Emerton)); Locket and Millidge (1953); Roberts (1987); Paquin and Dupérré (2003).

Records: M07 (2 males, 1 female). N = 3 adults; 2 males, 1 female.

Method: pitfall traps (2 males, 1 female).

Months: May (1 male); June (1 male); June-July (1 female).

Habitat: litter, Kalmia-Vaccinium heath (2 males, 1 female).

Regional Distribution: ME (Jennings et al., unpubl.); NY (Crosby and Bishop 1928, as Caseola 

alticeps Emerton)); NH, MA, QC (Buckle et al. 2001).

Pocadicnemis americana Millidge, 1976

Taxonomy: Millidge (1976); Paquin and Dupérré (2003).

Records: M07 (1 male, 2 females); M12 (2 males, 13 females); M24 (1 male); M26 (2 males). N = 

21 adults; 6 males, 15 females.

Methods: litter condenser-hand sorted (2 females); pitfall traps (6 males, 13 females).

Months: June (1 female); June-July (5 males); July (1 male, 2 females); July-August (1 female); 

August (11 females).
54

Habitats: sifted litter, Kalmia-Vaccinium heath (2 females); litter, Kalmia-Vaccinium heath (1 male); 

litter, freshwater marsh edge (2 males, 13 females); litter, red maple sapling stand (1 male); litter, 

saltmarsh edge (2 males).

Regional Distribution: ME (Hilburn and Jennings 1988); MA, CT, NY, ON, QC, NF (Buckle et 

al. 2001).

Satilatlas marxi Keyserling, 1886

Taxonomy: Millidge (1981b); Paquin and Dupérré (2003).

Records: M01 (1 male, 2 females); M06 (3 females); M07 (1 female). N = 7 adults; 1 male, 

6 females.

Methods: pitfall traps (1 male, 3 females); sweep net (3 females).

Months: May (4 females); June (1 male, 2 females).

Habitats: litter, seashore-backshore (1 male); litter, saltmarsh (2 females); litter, Kalmia-Vaccinium 

heath (1 female); sweeping grasses, saltmarshes (3 females).

Regional Distribution: ME (Millidge 1981b); MA, CT, RI, NY, ON, QC, NS (Buckle et al. 2001).

Sciastes truncatus (Emerton, 1882)

Taxonomy: Bishop and Crosby (1938); Kaston (1981); Millidge (1984); Paquin and Dupérré 

(2003).

Record: M54 (1 male). N = 1 adult male.

Method: litter condenser-Berlese funnel (1 male).

Month: September (1 male).

Habitat: sifted forest-fl oor litter, red spruce stand (1 male).

Regional Distribution: ME (Bishop and Crosby 1938); NH, VT, MA, NY, ON, QC, NB, NS, NF 

(Buckle et al. 2001). 

Scotinotylus exsectoides Millidge, 1981

Taxonomy: Millidge (1981c).

Records: M07 (1 female); M22 (1 female). N = 2 adult females.

Method: pitfall traps (2 females).

Months: May (1 female); July (1 female).
55

Habitats: litter, Kalmia-Vaccinium heath (1 female); litter, bigtooth aspen stand (1 female).

Regional Distribution: ME (Jennings et al., unpubl.); ON (Buckle et al. 2001).

Note: This species is known only by the female (Millidge 1981c); the male remains to be discovered 

and described.

Sisicottus montanus (Emerton, 1882)

Taxonomy: Bishop and Crosby (1938); Kaston (1981); Miller (1999); Paquin and Dupérré (2003).

Records: M01 (1 female); M04 (1 male, 2 females); M54 (1 female). N = 5 adults; 1 male, 4 females.

Methods: beating cloth (1 female); litter condenser-Berlese funnel (1 female); pitfall traps (1 male, 2 

females).

Months: May (1 female); May-June (1 female); June (1 female); July (1 male); September (1 female).

Habitats: beating red spruce foliage, old fi eld edge (1 female); litter, coastal red spruce stand (1 male, 

2 females); sifted forest-fl oor litter, red spruce stand (1 female).

Regional Distribution: ME (Emerton 1914, as Tmeticus montanus Emerton); NH, VT, MA, NY, 

ON, QC, NB, NF (Buckle et al. 2001).

Sisicus penifusifer Bishop & Crosby, 1938

Taxonomy: Bishop and Crosby (1938, as Sisicus penifusiferus Bishop & Crosby); Paquin and Dupérré 

(2003).

Record: M24 (1 female). N = 1 adult female.

Method: pitfall trap (1 female).

Month: May-June (1 female).

Habitat: litter, red maple sapling stand (1 female).

Regional Distribution: ME (Bishop and Crosby 1938); NY, ON, QC, NB (Buckle et al. 2001).

Soulgas corticarius (Emerton, 1909)

Taxonomy: Crosby and Bishop (1936); Kaston (1981); Paquin and Dupérré (2003).

Record: M01 (1 female). N = 1 adult female.

Method: search (1 female).

Month: October (1 female).
56

Habitat: on exterior wall, house (1 female).

Regional Distribution: ME (Jaros-Su et al., unpubl.); MA, CT, RI, NY, QC (Buckle et al. 2001).

Styloctetor stativus (Simon, 1881)

Taxonomy: Roberts (1987); Paquin and Dupérré (2003).

Record: M24 (1 male). N = 1 adult male.

Method: pitfall trap (1 male).

Month: June (1 male).

Habitat: litter, red maple sapling stand (1 male).

Regional Distribution: ME (Drummond et al., unpubl.); ON, QC (Buckle et al. 2001).

Tapinocyba simplex (Emerton, 1882)

Taxonomy: Crosby and Bishop (1933); Kaston (1981); Paquin and Dupérré (2003).

Records: M04 (1 female); M07 (1 female); M25 (2 females); M27 (1 female). N = 5 adult females.

Method: pitfall traps (5 females).

Months: May (1 female); May-June (3 females); July (1 female).

Habitats: litter, Kalmia-Vaccinium heath (1 female); litter, coastal red spruce stand (1 female); litter, 

white (paper) birch stand (2 females); litter, red maple-white (paper) birch stand (1 female).

Regional Distribution: ME (Crosby and Bishop 1933); NH, VT, MA, NY, ON, QC, NB, NF 

(Buckle et al. 2001).

*Tapinocyba sp.

Taxonomy: An unknown species per C. D. Dondale (pers. comm.).

Record: M01 (1 female). N = 1 adult female.

Method: search (1 female).

Month: October (1 female).

Habitat: “on my arm” (FG, Jr.).

Regional Distribution: ME (this study).

Note: This specimen is in the Canadian National Collection, Ottawa.
57

Tunagyna debilis (Banks, 1892)

Taxonomy: Kaston (1981); Millidge (1984); Paquin and Dupérré (2003).

Records: M04 (1 male); M06 (1 male); M24 (1 female). N = 3 adults; 2 males, 1 female.

Method: pitfall traps (2 males, 1 female).

Months: May-June (1 female); July (2 males).

Habitats: litter, saltmarsh (1 male); litter, coastal red spruce stand (1 male); litter, red maple sapling 

stand (1 female).

Regional Distribution: ME, (Millidge 1984); NH, MA, NY, ON, QC, NB, NS, NF (Buckle et al.

2001).

Walckenaeria atrotibialis O. Pickard-Cambridge, 1878

Taxonomy: Kaston (1981); Millidge (1983); Paquin and Dupérré (2003).

Records: M12 (2 males, 1 female); M26 (1 male). N = 4 adults; 3 males, 1 female.

Method: pitfall traps (3 males, 1 female).

Months: June-July (2 males); July-August (1 male); August (1 female).

Habitats: litter, freshwater marsh edge (2 males, 1 female); litter, saltmarsh edge (1 male).

Regional Distribution: ME (Millidge 1983); NH, VT, NY, ON, QC, NB, NS (Buckle et al. 2001).

Walckenaeria communis (Emerton, 1882)

Taxonomy: Kaston (1981); Millidge (1983); Paquin and Dupérré (2003).

Records: M01 (1 male); M04 (1 female); M12 (1 female). N = 3 adults; 1 male, 2 females.

Method: pitfall traps (1 male, 2 females).

Months: May-June (1 male); August (2 females).

Habitats: litter, deciduous-coniferous woodland (1 male); litter, coastal red spruce stand (1 female); 

litter, freshwater marsh edge (1 female).

Regional Distribution: ME (Jennings et al., unpubl.); NH, VT, MA, CT, RI, NY, QC, ON (Buckle 

et al. 2001).

Walckenaeria digitata (Emerton, 1913)

Taxonomy: Kaston (1981); Millidge (1983); Paquin and Dupérré (2003).
58

Records: M01 (1 female); M07 (1 male); M12 (1 female). N = 3 adults; 1 male, 2 females.

Method: pitfall traps (1 male, 2 females).

Months: June-July (1 male); July (1 female); August (1 female).

Habitats: litter, seashore-backshore (1 female); litter, Kalmia-Vaccinium heath (1 male); litter, 

freshwater pond edge (1 female).

Regional Distribution: ME (Jennings et al., unpubl.); NH, NY, ON, QC (Buckle et al. 2001).

Walckenaeria directa (O. Pickard-Cambridge, 1878)

Taxonomy: Kaston (1981); Millidge (1983); Paquin and Dupérré (2003).

Records: M04 (1 male, 2 females); M27 (1 male); M38 (1 female). N = 5 adults; 2 males, 3 females.

Methods: litter condenser-Berlese funnel (1 female); pitfall traps (2 males, 2 females).

Months: May (1 male); May-June (1 male); July (3 females).

Habitats: litter, coastal red spruce stand (1 male, 2 females); litter, red maple-white (paper) birch 

stand (1 male); sifted litter, Sphagnum-cranberry-rush bog (1 female).

Regional Distribution: ME (Crosby and Bishop 1931, Cornicularia directa Cambridge); NH, VT, 

MA, RI, CT, NY, ON, QC, NS (Buckle et al. 2001).

Walckenaeria exigua Millidge, 1983

Taxonomy: Millidge (1983); Paquin and Dupérré (2003).

Records: M04 (2 males, 3 females); M24 (1 male). N = 6 adults; 3 males, 3 females.

Methods: pitfall traps (3 males, 2 females); sifted litter-hand sorted (1 female).

Month: May (1 female); June (3 males); June-July (1 female); July (1 female).

Habitats: litter, coastal red spruce stand (2 males, 2 females); litter, red maple stand (1 male); sifted 

litter, coastal red spruce-maple stand (1 female).

Regional Distribution: ME (Millidge 1983); NH, MA, NY, ON, QC, NB, NS (Buckle et al. 2001).

Walckenaeria lepida (Kulczyński, 1885)

Taxonomy: Millidge (1983); Paquin and Dupérré (2003).

Records: M06 (1 female); M19 (1 female). N = 2 adult females.

Methods: beating cloth (1 female); pitfall trap (1 female).
59

Months: June (1 female); July (1 female).

Habitats: beating red spruce foliage, mixed hardwood-conifer (1 female); litter, saltmarsh (1 female).

Regional Distribution: ME (Jennings and Dimond 1988); NS (Buckle et al. 2001).

**Walckenaeria redneri Millidge, 1983

Taxonomy: Millidge (1983); Paquin and Dupérré (2003).

Record: M12 (1 female). N = 1 adult female.

Method: pitfall trap (1 female).

Month: July (1 female).

Habitat: litter, freshwater marsh edge (1 female).

Regional Distribution: ME (this study); QC (Buckle et al. 2001).

Walckenaeria spiralis (Emerton, 1882)

Taxonomy: Kaston (1981); Millidge (1983); Paquin and Dupérré (2003).

Record: M56 (1 female).

Method: litter condenser-hand sorted (1 female).

Month: September (1 female).

Habitat: sifted litter (grasses, forbs, tidal debris), salt meadow (1 female).

Regional Distribution: ME (Hilburn and Jennings 1988); MA, CT, RI, NY, ON, QC (Buckle et al.

2001).

Walckenaeria subspiralis Millidge, 1983

Taxonomy: Millidge (1983); Paquin and Dupérré (2003).

Records: M06 (1 female); M12 (1 female). N = 2 adult females.

Method: pitfall traps (2 females).

Months: May (1 female); August (1 female).

Habitats: litter, saltmarsh (1 female); litter, freshwater marsh edge (1 female).

Regional Distribution: ME (Jennings et al., unpubl.); NY, ON, QC, NB, NS (Buckle et al. 2001).
60

*Undetermined genus, species

Taxonomy: “Unknown” per C.D. Dondale (pers. comm.).

Record: M05 (1 female). N = 1 adult female.

Method: sweep net (1 female).

Month: May (1 female).

Habitat: sweeping wet meadow, old fi eld-brackish marsh edge (1 female).

Regional Distribution: ME (this study).

Note: This specimen is in the Canadian National Collection, Ottawa.
61

FAMILY TETRAGNATHIDAE

The tetragnathids or long-jawed orb weavers usually spin their webs near water. The common name, 

long-jawed orb weaver (also four-jawed spiders), stems from their large, powerful chelicerae, which 

in some species project forward. The web is sometimes large with many radii or small with few radii. 

The hub is bitten out after the spirals are in place (Dondale et al. 2003). Some species spin their 

webs over running water, others among tall grass of meadows and marshes, and others in shrubs and 

trees. Apparently juveniles of Pachygnatha spin webs, but the adults do not. Levi (2005b) provides 

additional information about family characteristics, natural history, and a key to the genera of North 

America north of Mexico.

Levi (2005b) also includes notes on the taxonomic history of this family. For example, Levi (1980b) 

lowered the Tetragnathidae to subfamily status; subsequently, Dondale et al. (2003) returned it to full 

family status. Early revisers included Meta and Zygiella in the Tetragnathidae or Araneidae; currently, 

Meta resides in the Tetragnathidae while Zygiella resides in the Araneidae. Important revisionary 

works, with keys to genera and species in the northern region, are given by Levi (1980b, 1981). The 

tetragnathid fauna in Maine is represented by 3 genera and 14 species; that of Milbridge by 3 genera 

and 10 species.

Meta ovalis (Gertsch, 1933)

Taxonomy: Levi (1980b, as Meta menardi (Latreille, 1804)); Kaston (1981, as Meta menardi 

(Latreille, 1804)); Dondale (1995); Paquin and Dupérré (2003).

Record: M01 (1 male). N = 1 adult male.

Method: search (1 male).

Month: August (1 male).

Habitat: on front door, house (1 male).

Regional Distribution: ME (Bryant 1908, as Meta menardi (Latreille) Simon); NH, VT, MA, CT, 

RI, NY, ON, QC, NS, NF (Levi 1980b, as Meta menardi (Latreille)).

Note: Gertsch (1933) fi rst described the species ovalis placing it in the genus Auchicybaeus, but later 

Gertsch and Ivie (1936) indicated that Auchicybaeus ovalis Gertsch, 1933 was a junior synonym 

of Meta menardi (Latreille, 1804), a European species. However, Marusik and Koponen (1992) 

concluded that the species in eastern North America was distinct from M. menardi, and named it M. 

americana Marusik & Koponen, 1992. Dondale (1995) showed that Gertsch’s earlier name ovalis has 

priority and that M. ovalis and M. americana represent a single species.

Pachygnatha autumnalis Marx, 1884

Taxonomy: Levi (1980b); Kaston (1981); Dondale et al. (2003); Paquin and Dupérré (2003).

Records: M01 (1 female); M07 (1 female). N = 2 adult females.
62

Methods: pitfall traps (2 females).

Months: June-July (1 female); August (1 female).

Habitats: litter, old fi eld edge and alder swamp (1 female); litter, Kalmia-Vaccinium heath (1 female).

Regional Distribution: ME (Procter 1946); NY (Crosby and Bishop 1928); CT (Kaston 1981); 

NH, VT, MA, ON (Levi 1980b); QC (Bélanger and Hutchinson 1992).

Pachygnatha brevis Keyserling, 1884

Taxonomy: Levi (1980b); Kaston (1981); Dondale et al. (2003); Paquin and Dupérré (2003).

Records: M01 5 males, 7 females); M05 (1 female); M06 (8 females); M17 (2 males, 3 females). N = 

26 adults; 7 males, 19 females.

Methods: pitfall trap (1 male); search (1 male); sweep net (5 males, 18 females); n. d. (1 female).

Months: May (3 males, 6 females); June (1 male, 1 female); August (3 males, 10 females); September 

(2 females).

Habitats: litter, seashore-backshore (1 male); sweeping grasses (Juncus-Spartina), saltmarshes (5 males, 

16 females); sweeping herbaceous vegetation, freshwater pond edge (2 females); at freshwater pond 

edge (1 male); n. d., old fi eld (1 female).

Regional Distribution: ME (Bishop 1923); NY (Crosby and Bishop 1928); CT (Kaston 1981); 

NH, VT, MA, RI, ON (Levi 1980b); QC (Bélanger and Hutchinson 1992).

Tetragnatha caudata Emerton, 1884

Taxonomy: Levi (1981); Kaston (1981); Dondale et al. (2003); Paquin and Dupérré (2003).

Records: M01 (1 male, 1 female); M12 (5 males, 9 females); M33 (4 males, 4 females); M55 (1 

male). N = 25 adults; 11 males, 14 females.

Method: sweep net (11 males, 14 females).

Months: June (8 males, 7 females); July (3 males, 7 females).

Habitats: sweeping grasses and other vegetation, freshwater pond edge (1 male, 1 female); sweeping 

marsh grasses, sedges, and shrubs, freshwater marshes (9 males, 13 females); sweeping grasses, 

freshwater streamside (1 male).

Regional Distribution: ME (Bryant 1908); NY (Crosby and Bishop 1928, as Eucta lacerta 

Walckenaer); MA, RI, ON (Levi 1981); QC (Bélanger and Hutchinson 1992).

Tetragnatha elongata Walckenaer, 1842

Taxonomy: Levi (1981); Kaston (1981); Dondale et al. (2003); Paquin and Dupérré (2003).
63

Records: M01 (3 males, 5 females). N = 8 adults; 3 males, 5 females.

Methods: search (2 males); sweep net (1 male, 5 females).

Months: June (2 males, 2 females); July (1 male); August (3 females).

Habitats: sweeping vegetation, freshwater pond edge (1 male, 5 females); on grasses, freshwater pond 

edge (1 male); on foundation, house (1 male).

Regional Distribution: ME (Bryant 1908); NY (Crosby and Bishop 1928); CT (Kaston 1981); MA, 

RI, ON, QC, NB (Levi 1981).

Tetragnatha extensa (Linnaeus, 1758)

Taxonomy: Levi (1981); Dondale et al. (2003); Paquin and Dupérré (2003).

Records: M01 (1 male); M06 (1 female); M17 (2 females); M33 (1 male, 1 female); M57 (1 male, 

4 females). N = 11 adults; 3 males, 8 females.

Method: sweep net (3 males, 8 females).

Months: June (2 males, 4 females); July (1 male, 1 female); August (3 females).

Habitats: sweeping vegetation, freshwater pond edge (1 male); sweeping marsh vegetation, 

saltmarshes (3 females); sweeping grasses and sedges, freshwater marsh (1 male, 1 female); sweeping 

rushes, brackish marsh (1 male, 4 females).

Regional Distribution: ME (Banks 1893); NY (Crosby and Bishop 1928); NH, ON, QC, NS, NF 

(Levi 1981).

*Tetragnatha guatemalensis O. Pickard-Cambridge, 1889

Taxonomy: Levi (1981); Kaston (1981); Dondale et al. (2003).

Records: M01 (2 females). N = 2 adult females.

Method: search (2 females).

Month: July (2 females).

Habitats: in shrubs on shore, seashore-backshore (1 female); on dead limb of Amelanchier sp., 

seashore-backshore (1 female).

Regional Distribution: ME (this study); CT (Kaston 1981); NY, ON, NS (Levi 1981).

Tetragnatha laboriosa Hentz, 1850

Taxonomy: Levi (1981); Kaston (1981); Dondale et al. (2003); Paquin and Dupérré (2003).
64

Records: M01 (5 males, 6 females); M05 (3 males); M06 (10 males, 3 females); M14 (1 female); 

M15 (1 female); M23 (2 females); M30 (1 female); M35 (2 females); M58 (6 males, 2 females). N = 

42 adults; 24 males, 18 females.

Methods: search (2 males, 4 females); sweep net (22 males, 13 females); n. d. (1 female).

Months: June (10 males, 5 females); July (11 males, 12 females); August (3 males, 1 female).

Habitats: sweeping vegetation, freshwater pond edge (1 male, 1 female); sweeping grasses on shore, 

seashore-backshore (2 males, 2 females); sweeping vegetation, old fi eld (2 males); sweeping vegetation 

(Juncus-Spartina), saltmarshes (11 males, 3 females); sweeping grasses and forbs, mixed hardwoodconifer (1 female); sweeping grasses and sedges, saltmarsh (1 female); sweeping roadside grasses, forbs, 

ferns, and rushes, alder-aspen edge (1 female); sweeping marsh grasses and rushes, brackish marsh (2 

females); sweeping grasses, forbs, and rushes, seashore-brackish marsh (6 males, 2 females); in web on 

meadowsweet bush, seashore-backshore (1 female); on grasses at shore, seashore-backshore (1 female); 

in high salt grass at shore, seashore-backshore (1 male); underside of wild Sarsaparilla leaf, old fi eld (1 

male); in old fi eld (1 female); at freshwater pond edge (Sphagnum-water lilies-sedges-grass), island (1 

female); on lawn chair (1 female).

Regional Distribution: ME (Bryant 1908); NY (Crosby and Bishop 1928); CT (Kaston 1981); NH, 

VT, MA, RI, ON, QC, NB, NS, NF (Levi 1981). 

Tetragnatha straminea Emerton, 1884

Taxonomy: Levi (1981); Kaston (1981); Dondale et al. (2003); Paquin and Dupérré (2003).

Records: M01 (1 male, 2 females); M14 (3 males, 4 females); M17 (1 male, 1 female); M24 (1 

female); M33 (2 females). N = 15 adults; 5 males, 10 females.

Methods: search (1 female); sweep net (5 males, 9 females).

Months: June (1 male, 3 females); July (3 males, 4 females); August (1 male, 2 females); September 

(1 female).

Habitats: sweeping grasses and sedges, seashore-backshore (1 female); sweeping vegetation, freshwater 

pond edge (1 male); sweeping understory vegetation (grasses, forbs) along trail, mixed hardwoodconifer (3 males, 4 females); sweeping grasses, saltmarsh (1 male, 1 female); sweeping understory 

vegetation, red maple sapling stand (1 female); sweeping grasses and sedges, freshwater marsh (1 

female); sweeping understory vegetation, freshwater marsh-mixed conifer-hardwood edge (1 female); 

on blackberry (1 female).

Regional Distribution: ME (Bryant 1908); NY (Crosby and Bishop 1928); CT (Kaston 1981); MA, 

ON (Levi 1981); QC (Bélanger and Hutchinson 1992).

Tetragnatha versicolor Walckenaer, 1842

Taxonomy: Levi (1981); Kaston (1981); Dondale et al. (2003); Paquin and Dupérré (2003).
65

Records: M01 (5 males, 8 females); M04 (1 male); M06 (2 females); M12 (4 females); M14 (1 

male); M17 (1 female); M23 (1 male); M47 (1 female); M49 (1 female). N = 25 adults; 8 males, 17 

females.

Methods: beating cloth (3 females); search (1 male, 2 females); shake-sweep net (1 female); sweep net 

(6 males, 11 females); n. d. (1 male).

Months: May (1 male, 1 female); June (1 male, 4 females); July (4 males, 7 females); August (2 

males, 5 females).

Habitats: beating alder foliage (2 females); beating red spruce foliage, seashore-backshore (1 female); 

shaking red spruce foliage over sweep net, saltmarsh edge (1 female); sweeping vegetation, freshwater 

pond edge (4 males, 4 females); sweeping vegetation, old fi eld (1 female); sweeping vegetation 

near brackish pond, seashore-brackish marsh (1 male); sweeping vegetation, saltmarsh (2 females); 

sweeping marsh grasses, sedges, and meadowsweet freshwater marsh edge (4 females); sweeping 

understory vegetation along trail, mixed hardwood-conifer (1 male); sweeping marsh vegetation, 

freshwater streamside (1 female); n. d., spruce, mixed conifer-hardwood, island (1 male); on low 

understory foliage, Betula-Acer stand (1 male); on wall, shed (1 female).

Regional Distribution: ME (Seeley 1928, as Tetragnatha extensa); CT (Kaston 1981); NH, VT, NY, 

MA, RI, ON, QC, NB, NS, NF (Levi 1981).

Note: Levi (1981) indicates that Tetragnatha extensa of Seeley (1928) is T. versicolor Walckenaer, 

1842.
66

FAMILY ARANEIDAE

The araneids or orb weavers include numerous genera and species; in species richness, they are the 

third largest spider family in the world (Dondale et al. 2003). This family includes large-bodied 

spiders whose webs are often seen in gardens, parks, meadows, old fi elds, and forests. Many spin large 

orb webs that are strikingly beautiful when covered with dew. Some spin retreats in folded leaves, but 

usually have an attachment line or “signal line” to the nearby orb web. Others remain at the center 

of the web in a “head-down” position. Some species add a zigzag pattern of silk, the stabilimentum, 

above and below the web’s center.

Several morphological characters are unique to at least one representative of the Araneidae, but 

none are diagnostic for all (Dondale et al. 2003). These characters include protruding anterior 

median eyes; lateral eyes together or on tubercles; abdomen with or without dorsal humps; abdomen 

with or without lateral spines; and presence or absence of coxal hook on leg I of males. Additional 

character descriptors and notes on natural and taxonomic histories are given by Levi (2005a). Levi 

(2002, 2005a) provides identifi cation keys to the theridiid genera of North America. In Maine, the 

Araneidae are represented by 16 genera and 44 species; in Milbridge, by 12 genera and 24 species.

Acanthepeira stellata (Walckenaer, 1805)

Taxonomy: Levi (1976); Kaston (1981); Dondale et al. (2003); Paquin and Dupérré (2003).

Record: M55 (1 female). N = 1 adult female.

Method: sweep net (1 female).

Month: June (1 female).

Habitat: sweeping grasses and forbs, playground edge (1 female).

Regional Distribution: ME (Procter 1933, as Marxia stellata (Walckenaer); NH, VT, MA, CT, RI, 

NY, ON, QC (Levi 1976).

Araneus cavaticus (Keyserling, 1882)

Taxonomy: Levi (1971); Kaston (1981); Dondale et al. (2003); Paquin and Dupérré (2003).

Records: M01 (1 male, 1 female); M50 (1 female). N = 3 adults; 1 male, 2 females.

Method: search (1 male, 2 females).

Month: August (1 male, 2 females).

Habitats: in orb web, camp house (1 female); in orb web, house porch ceiling (1 female); on screen 

door below kiwi vine, house (1 male).

Regional Distribution: ME (Bryant 1908, as Araneus cavaticus (Keyserling) Simon); NH, VT, MA, 

CT, RI, NY, ON, QC, NB (Levi 1971).
67

Araneus iviei (Archer, 1951)

Taxonomy: Levi (1971); Dondale et al. (2003); Paquin and Dupérré (2003).

Record: M01 (1 male). N = 1 adult male.

Method: search (1 male).

Month: July (1 male).

Habitat: on dead stem, meadowsweet (1 male).

Regional Distribution: ME, VT, MA, CT, NY (Levi 1971); QC Paquin and Dupérré (2003).

Araneus juniperi (Emerton, 1884)

Taxonomy: Levi (1973); Kaston (1981); Dondale et al. (2003); Paquin and Dupérré (2003).

Record: M35 (1 male). N = 1 adult male.

Method: sweep net (1 male).

Month: July (1 male).

Habitat: sweeping brackish marsh (1 male).

Regional Distribution: ME (Emerton 1884, as Epeira juniperi Emerton); NH, MA, NY (Levi 

1973); CT (Kaston 1981); QC (Paquin and Dupérré 2003).

Araneus marmoreus Clerck, 1757

Taxonomy: Levi (1971); Kaston (1981); Dondale et al. (2003); Paquin and Dupérré (2003).

Records: M01 (11 males, 37 females); M04 (2 females); M05 (1 female); M06 (5 females); M12 (1 

female); M13 (2 females); M18 (1 female); M22 (1 female); M29 (1 female); M38 (1 male); M47 (1 

female); M48 (2 females); M56 (1 female). N = 67 adults; 12 males, 55 females.

Methods: beating cloth (1 female); brushing tree-bole bark (2 females); pitfall trap (1 female); search 

(11 males, 44 females); sweep net (1 male, 5 females); n. d. (2 females).

Months: July (1 male, 1 female); August (11 males, 30 females); September (22 females); October (2 

females).

Habitats: in folded leaf retreats, over webs in blackberry shrubs (3 females); in folded red maple 

leaves, over orb webs (2 females); in folded leaf retreats, over webs in alder (2 females); in dome 

(retreat) among alder leaves, over orb web (1 female); in folded leaf over web between two small 

alders, old fi eld edge (1 female); in folded leaf retreat above web among aster and Amelanchier sp. (1 

female); in retreat above web on white (paper) birch, seashore-backshore (1 female); in folded leaf, 
68

small pin cherry above orb web (1 female); in retreat above web, small wild cherry shrubs, old fi eld 

edge (2 females); in rolled aster leaf over orb web, old fi eld (1 female); in rolled raspberry leaf over 

orb web (1 female); in folded spruce (Picea sp.) at seashore (1 female); in leaf retreat, gray birch, 

Betula-Acer stand (1 female); in dome (retreat) among Amelanchier leaves (1 female); in retreat on 

meadowsweet, old fi eld (1 female); in cone-shaped retreat of leaves, old fi eld (1 female); in wild 

blackberry (1 male); in retreat on roadside chokecherry, mixed hardwood-conifer (1 female); in 

retreat above orb web, roadside northern red oak (1 female); in retreat above web, roadside Juncus sp., 

mixed conifer (1 female); in web on red spruce, near seaside causeway (1 female); in orb web between 

goldenrod and fern, old fi eld (1 female); in center of web among goldenrod, old fi eld (1 female); 

in web, granite cliff face, seashore (1 female); in alder, freshwater marsh edge (1 female); in web, 

saltmarsh (2 females); in web, tall grass, saltmarsh (1 female); in web, n. d. (1 female); among tall 

weeds, saltmarsh (1 female); on trunk of white (paper) birch, Betula-Acer stand (1 female); on apple 

tree ascending dragline (1 male); on goldenrod, old fi eld (1 male); on potato plants, vegetable garden 

(1 female); on underside of blackberry leaf below agelenid web, old fi eld edge (1 male); beating 

foliage in alder thicket (1 female); brushing bark of white (paper) birch, Betula-Acer stand (2 females); 

sweeping vegetation of old fi eld (2 females); sweeping vegetation, old fi eld edge (1 female); sweeping 

young alders, roadside (1 female); sweeping Sphagnum-cranberry-bog (1 male); sweeping grasses 

and seaside goldenrod, salt meadow (1 female); litter, bigtooth aspen stand (1 female); in retreat 

above web, boat (1 female); in orb web on house (1 female); in retreat above orb web, house porch 

(1 female); in retreat inside lamp cover, shingled house wall (1 female); in folded patio umbrella (1 

male); in web under bridge, freshwater stream (1 female); on side of house (2 males,1 female); on 

mailbox, roadside (1 female); on dragline, house exterior (1 male); on porch fl oor, house (1 male); on 

foliage in garden, no web (1 male); on lawn chair (1 male); alder, seashore-backshore (1 female); old 

fi eld (1 female); n. d. (1 female).

Regional Distribution: ME (Bryant 1908, as Araneus marmoreus Fabricius); NH, VT, MA, CT, RI, 

NY, ON, QC, NB, NS (Levi 1971).

Araneus nordmanni (Thorell, 1870)

Taxonomy: Levi (1971); Kaston (1981); Dondale et al. (2003); Paquin and Dupérré (2003).

Records: M01 (2 females); M04 (1 male, 1 female); M05 (1 female); M06 (1 female); M08 (2 

females); M54 (1 female). N = 9 adults; 1 male, 8 females.

Methods: search (1 male, 6 females); sweep net (1 female); n. d. (1 female).

Months: August (1 male, 4 females); September (4 females).

Habitats: in orb webs, roadside shrubs (2 females); in web, small balsam fi r, seaside (1 female); 

in web, saltmarsh (1 female); in web on red spruce, old fi eld edge (1 female); under loose bark of 

white (paper) birch, mixed conifer-hardwood (1 male); sweeping plants, coastal red spruce-mixed 

hardwoods (1 female); old fi eld-brackish marsh (1 female); in web on side of shed (1 female).

Regional Distribution: ME (Emerton 1884, as Epeira nordmanni Thorell); NH, VT, MA, RI, NY, 

ON, QC, NB, NS (Levi 1971).
69

Araneus pratensis (Emerton, 1884)

Taxonomy: Levi (1973); Kaston (1981); Dondale et al. (2003).

Records: M01 (2 males); M07 (1 female). N = 3 adults; 2 males, 1 female.

Methods: sweep net (2 males); unsifted litter-Berlese funnel (1 female).

Months: June (2 males); August (1 female).

Habitats: sweeping grasses, forbs, and shrubs, old fi eld (2 males); litter, Kalmia-Vaccinium heath.

Regional Distribution: ME (Procter 1946, as Singa pratensis Emerton); NH, VT, MA, CT, NY 

(Levi 1973); ON (Levi 1975a).

Araneus saevus (L. Koch, 1872)

Taxonomy: Levi (1971); Kaston (1981); Dondale et al. (2003); Paquin and Dupérré (2003).

Records: M01 (4 males). N = 4 adult males.

Method: search (4 males).

Months: September (1 male); October (3 males).

Habitats: on lilac leaf (1 male); walking across porch, house (1 male); on outside wall of house (1 

male); on outside of front door, house (1 male).

Regional Distribution: ME (Procter 1938, as Aranea solitaria (Emerton); NH, VT, NY, ON, QC, 

NB, NS, NF (Levi 1971).

Araneus trifolium (Hentz, 1847)

Taxonomy: Levi (1971); Kaston (1981); Dondale et al. (2003); Paquin and Dupérré (2003).

Records: M01 (3 males, 10 females); M05 (3 males, 1 female); M06 (4 males, 4 females); M38 

(1 female); M56 (1 female). N = 27 adults; 10 males, 17 females.

Methods: search (3 males, 11 females); sweep net (7 males, 6 females).

Months: July (1 male, 1 female); August (9 males, 11 females); September (5 females).

Habitats: sweeping vegetation of old fi elds (3 males, 3 females); sweeping grasses, saltmarsh (4 males, 

2 females); sweeping grasses and seaside goldenrod, salt meadow (1 female); in rolled leaf over web, 

seaside goldenrod, saltmarsh (1 female); in folded leaf retreat, ornamental cinquefoil (1 female); in 

folded leaf (Spiraea sp.) retreat, old fi eld (1 female); in folded leaf over orb web, old fi eld (1 male); 

in retreat, Juncus sp. seedhead, Sphagnum-cranberry-rush bog (1 female); in vegetation of old fi eld 
70

(4 females); in orb web with prey, fl ower garden (1 female); under leaf near web, saltmarsh (1 female); 

on lily in garden (1 female); on automobile door (1 male); on outside wall of house (1 male).

Regional Distribution: ME (Hentz 1875, as Epeira trifolium); NH, VT, MA, CT, RI, NY, ON, QC, 

NB, NS, NF, LB (Levi 1971).

Araniella displicata (Hentz, 1847)

Taxonomy: Levi (1974); Kaston (1981); Dondale et al. (2003); Paquin and Dupérré (2003).

Records: M01 (9 males, 13 females); M12 (1 male, 1 female); M14 (1 female); M23 (1 female); M35 

(1 female); M51 (1 male, 2 females). N = 30 adults; 11 males, 19 females.

Methods: beating cloth (1 male, 2 females); pitfall trap (1 male); search (5 males, 12 females); sweep 

net (4 males, 4 females); shake-sweep net (1 female).

Months: May (2 females); June (8 males, 7 females); July (3 males, 9 females); August (1 female).

Habitats: beating red spruce foliage, old fi eld edge (1 female); beating red spruce foliage, mixed 

conifer (1 male, 1 female); sweeping vegetation, old fi eld (1 male, 1 female); sweeping vegetation, old 

fi eld edge (1 male); sweeping Vaccinium and Myrica near woods (1 female); sweeping low vegetation, 

deciduous-coniferous woodland (1 male); sweeping grasses on shore, seashore (1 male); sweeping 

grasses, brackish marsh (1 female); sweeping grasses and Kalmia, mixed conifer opening, island (1 

female); shaking speckled alder bushes over sweep net, mixed conifer opening (1 female); in small 

web on mountain-ash, with prey (1 female); in web on wild cherry leaf, along lane (1 female); in 

shrubbery (1 male, 1 female); in alder thicket (1 female); in webs on blackberry leaves, old fi eld (3 

females); in web, top of alder leaf, along lane (1 female); in web, garden (1 male); in web along trail, 

mixed hardwood-conifer understory (1 female); on willow shrub leaf, freshwater pond edge (1 male); 

on willow shrub leaf, freshwater marsh edge (1 female); litter, freshwater marsh edge (1 male); in shed 

(1 male); on side of house wall (1 female); on bedroom wall, house (1 male); on bedroom ceiling, 

house (1 female).

Regional Distribution: ME (Bryant 1908, as Araneus displicatus (Hentz) Simon); NH, VT, MA, CT, 

RI, NY, ON, QC, NS, NF, LB (Levi 1974).

Argiope aurantia (Lucas, 1833)

Taxonomy: Levi (1968); Kaston (1981); Dondale et al. (2003); Paquin and Dupérré (2003).

Records: M01 (7 females); M05 (2 females); M06 (2 males, 3 females); M18 (1 female); M21 (1 

female); M38 (1 female); M56 (1 female). N = 18 adults; 2 males, 16 females.

Methods: search (8 females); sweep net (2 males, 6 females); n. d. (2 females).

Months: August (2 males, 8 females); September (4 females); October (4 females).

Habitats: sweeping saltmarsh vegetation (2 males, 1 female); sweeping old fi eld vegetation (3 

females); sweeping Sphagnum-cranberry-rush bog (1 female); sweeping grasses and seaside goldenrod, 
71

salt meadow (1 female); in web among low forbs, old fi eld (1 female); in web with wrapped prey, old 

fi eld (1 female); among tall weeds, saltmarsh (1 female); in orb web, roadside goldenrod (1 female); 

in web, vegetable garden (1 female); on and under bark of dead tree, mixed hardwood-conifer (1 

female); in window, house (1 female); in storage cabinet, Refuge headquarters building (1 female); n. 

d. (2 females).

Regional Distribution: ME (Bryant 1908); NH, MA, CT, RI, NY, ON, NS (Levi 1968); QC 

(Bélanger and Hutchinson 1992).

Argiope trifasciata (Forskål, 1775)

Taxonomy: Levi (1968); Kaston (1981); Dondale et al. (2003); Paquin and Dupérré (2003).

Records: M01 (4 males, 9 females); M05 (23 males, 5 females); M06 (13 males, 3 females); M13 (4 

males); M21 (1 male); M38 (9 males, 5 females); M56 (8 males, 5 females). N = 89 adults; 62 males, 

27 females.

Methods: search (5 males, 2 females); sweep net (50 males, 23 females); n. d. (7 males, 2 females). 

Months: August (50 males, 14 females); September (11 males, 11 females); October (2 females); n. d. 

(1 male).

Habitats: sweeping grasses, forbs, shrubs, small trees, old fi eld (2 males, 3 females); sweeping old 

fi elds (13 males, 6 females); sweeping aster, goldenrod, and small alders, old fi eld (1 female); n. d., 

old fi eld (7 males, 2 females); sweeping grasses and other vegetation, saltmarsh (12 males, 3 females); 

sweeping Sphagnum-cranberry-rush bog (9 males, 5 females); sweeping grasses and forbs, including 

seaside goldenrod, salt meadow (8 males, 5 females); sweeping brackish marsh near river (4 males); 

sweeping low foliage, freshwater streamside (4 males); among tall weeds near female, saltmarsh (1 

male); in center of web, old fi eld (2 females); in shrubs, old fi eld (1 male); on truck fender, Refuge 

headquarters (1 male).

Regional Distribution: ME (Emerton 1894, as Argiope transversa Emerton); NH, VT, MA, CT, RI, 

NY, ON, QC, NS (Levi 1968)

Cyclosa conica (Pallas, 1772)

Taxonomy: Levi (1977); Kaston (1981); Dondale et al. (2003); Paquin and Dupérré (2003).

Records: M01 (1 male, 1 female); M04 (2 females); M23 (1 female); M35 (1 male); M49 (1 female). 

N = 7 adults; 2 males, 5 females.

Methods: beating cloth (1 female); search (1 male, 1 female); sweep net (1 male, 1 female); n. d. 

(2 females).

Months: June (1 male, 1 female); July (1 male, 3 females); August (1 female).

Habitats: beating red spruce foliage, seashore-backshore (1 female); sweeping grasses, forbs, and 

shrubs, roadside-red spruce forest (1 female); sweeping marsh grasses and rushes, brackish marsh 
72

(1 male); red spruce, Sphagnum moss, island (1 female); in center of orb web, Betula-Acer stand 

(1 female); on side of shed (1 male); n. d. (1 female).

Regional Distribution: ME (Emerton 1884); NH, VT, MA, CT, NY, ON, QC, NB, NS, NF, LB 

(Levi 1977).

Eustala anastera (Walckenaer, 1842)

Taxonomy: Levi (1977); Kaston (1981); Dondale et al. (2003); Paquin and Dupérré (2003).

Records: M01 (1 female); M51 (1 male). N = 2 adults; 1 male, 1 female.

Method: sweep net (1 male, 1 female).

Months: June (1 male); July (1 female).

Habitats: sweeping rhodora and Spiraea, mixed conifer opening (1 male); sweeping grasses along 

saltwater creek (1 female).

Regional Distribution: ME, NH, MA, CT, RI, NY, ON, QC, NB, NS (Levi 1977).

*Eustala cepina (Walckenaer, 1842)

Taxonomy: Levi (1977); Dondale et al. (2003).

Record: M55 (1 female). N = 1 adult female.

Method: sweep net (1 female).

Month: June (1 female).

Habitat: sweeping grasses and forbs, edge of school playground-disturbed area (1 female).

Regional Distribution: ME (this study); MA, CT, NY, ON (Levi 1977); NS (Dondale et al. (2003).

Hypsosinga funebris (Keyserling, 1892)

Taxonomy: Levi (1972, as Hypsosinga singaeformis (Scheffer, 1904)); Levi (1975a); Dondale et al.

(2003).

Records: M01 (1 male); M04 (1 female). N = 2 adults; 1 male, 1 female.

Methods: search (1 male); sweep net (1 female).

Month: June (1 male, 1 female).

Habitats: sweeping shrubbery vegetation, coastal mixed conifer (1 female); on wall of house (1 male).
73

Regional Distribution: ME, NH, VT, MA, CT, NY (Levi 1972); ON (Dondale et al. 2003).

Hypsosinga pygmaea (Sundevall, 1831)

Taxonomy: Levi (1972, as Hypsosinga variabilis (Emerton, 1884); Levi (1975a); Kaston (1981); 

Dondale et al. (2003); Paquin and Dupérré (2003).

Records: M05 (3 females); M06 (26 females); M12 (4 females); M33 (1 female); M47 (2 females); 

M55 (4 females); M57 (7 females); M58 (7 females). N = 54 adult females.

Method: sweep net (54 females).

Months: May (9 females); June (43 females); July (2 females).

Habitats: sweeping vegetation (Juncus-Spartina), saltmarsh (26 females); sweeping rushes (Juncus sp.), 

brackish marsh (7 females); sweeping marsh vegetation near brackish pond, brackish marsh-seashore 

(7 females); sweeping marsh grasses and shrubs (Spiraea-Kalmia), freshwater marshes (5 females); 

sweeping grasses and forbs, edge of playground (3 females); sweeping grasses along freshwater streams 

(3 females); sweeping wet meadow, wet meadow-brackish marsh (2 females); sweeping grasses and 

forbs, old fi eld (1 female).

Regional Distribution: ME (Emerton 1884, as Singa variabilis Emerton); NH, VT, MA, CT, NY, 

ON, QC, LB (Levi 1972); NS (Levi 1975a); RI, NB, NF (Dondale et al. 2003).

Hypsosinga rubens (Hentz, 1847)

Taxonomy: Levi (1972); Kaston (1981); Dondale et al. (2003); Paquin and Dupérré (2003).

Record: M32 (1 female). N = 1 adult female.

Method: beating cloth (1 female).

Month: July (1 female).

Habitat: beating balsam fi r foliage, mixed hardwood-conifer (1 female).

Regional Distribution: ME (Sferra et al., unpubl.); NH, MA, CT, RI, NY, ON, NS (Levi 1972); 

QC (Bélanger and Hutchinson 1992); NB, NF (Dondale et al. 2003).

Larinioides cornutus (Clerck, 1757)

Taxonomy: Levi (1974, as Nuctenea cornuta (Clerck)); Kaston (1981, as Nuctenea cornuta (Clerck)); 

Dondale et al. (2003); Paquin and Dupérré (2003).

Records: M01 (1 female); M12 (1 female); M14 (1 male); M33 (1 female). N = 4 adults; 1 male, 

3 females.

Methods: search (1 female); sweep net (1 male, 2 females).
74

Months: June (2 females); August (1 male, 1 female).

Habitats: sweeping grasses and sedges, freshwater marsh (1 male, 1 female); sweeping vegetation, old 

fi eld (1 female); in silk-tied retreat, blue-joint grass seedhead, freshwater marsh (1 female).

Regional Distribution: ME (Hentz 1875, as Epeira strix); NH, VT, MA, CT, RI, NY, ON, QC, NB, 

NS, NF, LB (Levi 1974; Dondale et al. 2003).

Larinioides patagiatus (Clerck, 1757)

Taxonomy: Levi (1974, as Nuctenea patagiata (Clerck)); Kaston (1981, as Nuctenea patagiata 

(Clerck)); Dondale et al. (2003); Paquin and Dupérré (2003).

Records: M01 (1 male, 3 females); M30 (1 female). N = 5 adults; 1 male, 4 females.

Method: search (1 male, 4 females).

Months: June (1 male, 1 female); July (2 females); August (1 female).

Habitats: in retreat, rolled alder leaf, balsam fi r-red spruce stand (1 female); dropping from dead 

branch of Amelanchier sp., riparian (1 female); in rural mailbox, roadside (1 male, 2 females).

Regional Distribution: ME (Emerton 1884, as Epeira patagiata (Clerck)); NH, VT, MA, CT, RI, 

NY, ON, QC, NB, NS, NF, LB (Levi 1974, Dondale et al. 2003).

Mangora placida (Hentz, 1847)

Taxonomy: Levi (1975b); Kaston (1981); Dondale et al. (2003); Paquin and Dupérré (2003).

Records: M01 (5 females); M04 (1 female); M10 (2 females); M25 (5 females). N = 13 adult females.

Methods: beating cloth (2 females); search (3 females); sweep net (8 males).

Months: May (2 females); June (10 females); July (1 female).

Habitats: sweeping understory vegetation, white (paper) birch stand (5 females); sweeping vegetation, 

freshwater pond edge (2 females); sweeping grasses, forbs, and shrubs, roadside, red spruce forest (1 

female); beating red spruce foliage (saplings), conifer-mixed hardwood (2 females); in alder (1 female); 

in web on small quaking aspen (1 female); under cardboard, Amelanchier-Rubus stand (1 female).

Regional Distribution: ME (Bryant 1908, as Mangora placida (Hentz) Simon); NH, MA, CT, RI, 

NY, ON, QC (Levi 1975b); VT, NB, NS (Dondale et al. 2003).

Metepeira palustris Chamberlin & Ivie, 1942

Taxonomy: Levi (1977, as Metepeira grandiosa palustris Chamberlin & Ivie); Dondale et al. (2003); 

Paquin and Dupérré (2003).
75

Records: M01 (2 males, 6 females); M06 (1 female). N = 9 adults; 2 males, 7 females.

Methods: search (2 males, 6 females); sweep net (1 female).

Months: August (2 males, 6 females); September (1 female).

Habitats: in composite webs, low shrubs (1 male, 2 females); in web on small trees and shrubs, ledge, 

mixed hardwood conifer (1 female); in silk debris hung from marsh grasses, saltmarsh (1 female); in 

old fi eld (1 female); sweeping old fi eld (1 female); on aster, old fi eld (1 female); on rural mailbox, 

roadside (1 male).

Regional Distribution: ME (Emerton 1915, as Epeira labyrinthea Hentz); NY, ON, QC, NS (Levi 

1977, Dondale et al. 2003).

Note: Levi (1977) indicated that Emerton’s “bog variety” of Epeira labyrinthea was a junior synonym 

of Metepeira grandiosa Chamberlin & Ivie, 1941, and assigned this variety to one of three subspecies, 

i.e., M. grandiosa palustris. However, Dondale et al. (2003) elevated M. palustris to full species rank. 

Neoscona arabesca (Walckenaer, 1842)

Taxonomy: Berman and Levi (1971); Kaston (1981); Dondale et al. (2003); Paquin and Dupérré 

(2003).

Records: M01 (6 males, 37 females); M02 (1 male); M04 (3 males, 6 females); M05 (5 males, 15 

females); M06 (14 males, 24 females); M07 (4 males, 1 female); M12 (1 male, 3 females); M17 (1 

female); M23 (1 male, 1 female); M28 (1 male); M29 (1 male); M30 (1 male); M34 (1 male); M35 

(1 male); M38 (4 males, 3 females); M47 (11 males, 2 females); M48 (1 female); M55 (1 male, 1 

female); M56 (5 females). N = 156 adults; 56 males, 100 females.

Methods: beating cloth (4 males, 1 female); pitfall traps (1 male, 1 female); search (3 males, 32 

females); sweep net (44 males, 61 females); n. d. (4 males, 5 females).

Months: June (1 male, 1 female): July (8 males, 10 females); August (47 males, 70 females); 

September (19 females).

Habitats: sweeping old fi elds (8 males, 15 females); sweeping saltmarshes (12 males, 24 females); 

sweeping grasses and other marsh vegetation near freshwater stream, riparian (11 males, 2 females); 

sweeping Sphagnum-cranberry-rush bog (4 males, 3 females); sweeping grasses and seaside goldenrod, 

salt meadow (5 females); sweeping brackish marsh-old fi eld near river (1 male, 4 females); sweeping 

grasses along saltwater creek (1 male, 3 females); sweeping vegetation in low marshy area, coastal 

spruce-mixed hardwoods (1 male, 2 females); sweeping freshwater marsh (3 females); sweeping 

Kalmia-Vaccinium heath (2 males); sweeping grasses and forbs, playground edge (1 male, 1 female); 

sweeping white and yellow clover, roadside disturbed area (1 male); sweeping grasses, forbs, ferns, 

and rushes, roadside alder-aspen edge (1 male); sweeping grasses, sedges, and forbs, dry streambed, 

mixed hardwood-conifer (1 male); sweeping brackish marsh, inland (1 male); beating red spruce 

foliage, mixed conifer (1 male, 1 female); beating red spruce, roadside (1 male); beating eastern larch 

foliage, mixed conifer (1 male); beating spruce foliage, seashore-backshore (1 male); litter, saltmarsh 
76

(1 female); in saltmarsh (2 males); litter, freshwater marsh edge (1 male); in webs, on foliage, on 

zucchini, and running on ground, vegetable garden (4 females); in orb webs among goldenrod, in old 

fi eld (4 females); in orb webs on Amelanchier sp. (2 females); in web, old fi eld (1 female); in shrubs, 

old fi eld (1 female); in fl ower garden (1 female); in retreat above web on Spiraea and timothy grass, 

old fi eld (1 female); climbing on timothy grass, old fi eld (1 female); in center of orb web on barberry 

shrub (1 female); in center of orb web on red spruce (1 female); in center of orb web, n. d. (1 female); 

in orb web among alders (1 female); in web, small trees and shrubs on ledge, mixed hardwood-conifer 

(1 female); in low plants, edge of lawn (1 female); in orb web, dining room wall, house (1 female); in 

center of orb web, house window (1 female); in web on side of house (1 female); on collector’s pants, 

old fi eld (1 female); on fl ower head of Helianthus sp., fl ower garden (1 female); on foliage, freshwater 

pond edge (1 female); on patio, house (1 female); on screen door, house (1 female); on side of house 

(1 male); on cobblestone beach, seashore (1 female); under bark of dead red maple, mixed hardwoodconifer (1 female); on heath plants, Kalmia-Vaccinium heath (2 males, 1 female); in web on shore 

ledge, seashore (1 female); n. d. (1 male, 3 females).

Regional Distribution: ME (Bryant 1908, as Araneus trivittatus (Keyserling) Comstock); NH, VT, 

MA, CT, RI, NY, ON, QC, NB, NS (Berman and Levi 1971, Dondale et al. 2003).

Note: During outbreaks of the spruce budworm, this spider captures and feeds on budworm moths 

entangled in its web (Jennings and Houseweart 1988).

Zygiella atrica (C. Koch, 1845)

Taxonomy: Gertsch (1964); Levi (1974); Kaston (1981); Paquin and Dupérré (2003).

Records: M04 (1 male, 1 female); M21 (1 male, 2 females). N = 5 adults; 2 males, 3 females.

Methods: beating cloth (1 male, 1 female); search (1 male, 2 females).

Months: August (1 male, 1 female); September (1 male, 2 females).

Habitats: beating red spruce foliage, seashore-backshore (1 male, 1 female); lumber pile near Refuge 

headquarters building (1 male, 2 females).

Regional Distribution: ME (Emerton 1914); MA, RI, NY, ON (Gertsch 1964); NS (Levi 1974); 

QC (Bélanger and Hutchinson 1992); NF (Dondale et al. 2003).

Note: This is an introduced Palearctic species in North America (Dondale et al. 2003) that, according 

to Gertsch (1964), was fi rst noticed by H. C. McCook at Annisquam, MA in about 1885.
77

FAMILY AGELENIDAE

The agelenids or funnel-web weavers spin sheet or platform webs with a tube or funnel leading off 

one edge. Several strands of silk are spun above the sheet, which serve as barriers to fl ying insects. 

Insects that encounter these upward strands may fall to the sheet below and become potential prey for 

the resident spider, which usually hides in the tube or funnel. If potential prey is perceived, the spider 

rushes out of its retreat and infl icts one or more bites. Once the prey is subdued, the spider seizes it 

and returns to the funnel to feed.

Some agelenids resemble lycosids or wolf spiders; however, the eyes of agelenids are arranged in two 

rows, not three as in the Lycosidae. The spinnerets also project posteriorly and consist of longer 

segments than lycosid spinnerets. The tarsi bear three claws but lack scopulae. Other characters of 

the family Agelenidae are described by Bennett and Ubick (2005), who also provide notes on natural 

history and a key to the genera of North America. 

Several genera formerly assigned to the Agelenidae have been removed and assigned to other families. 

For example, Cicurina now belongs to the Dictynidae; Coras and Wadotes are now considered 

members of the Amaurobiidae. With these changes, the agelenid fauna of Maine now consists of only 

two genera and seven species; both genera and four species have been found in Milbridge. 

Agelenopsis actuosa (Gertsch & Ivie, 1936)

Taxonomy: Chamberlin and Ivie (1941); Roth and Brown (1986); Paquin and Dupérré (2003).

Records: M01 (2 males, 3 females); M04 (3 males, 3 females); M05 (1 male, 4 females); M06 (3 

males, 3 females); M07 (1 female); M13 (6 females); M38 (3 females); M39 (1 female). N = 33 

adults; 9 males, 24 females.

Methods: pitfall traps (4 males, 1 female); search (4 males, 14 females); sweep net (7 females); n. d. 

(1 male, 2 females).

Months: July (1 female); July-August (1 male); August (7 males, 23 females); August-September (1 

male).

Habitats: in funnel webs, roadside, mixed conifer-hardwood (6 females); in funnel webs, old fi eld 

(1 male, 2 females); in funnel webs, upper beach, seashore (2 females); in funnel webs, among rocks, 

coastal spruce-mixed hardwood (2 males); in webbed conical retreat on blackberry (Rubus sp.) (1 

male, 1 female); in web, saltmarsh (1 female); litter, saltmarsh (3 males); litter, seashore-backshore 

(1 male); litter, Kalmia-Vaccinium heath (1 female); sweeping saltmarsh (1 female); sweeping 

Sphagnum-cranberry-rush bog (3 females); sweeping old fi eld (2 females); sweeping understory shrubs 

and ferns, red maple stand (1 female); in low plants, edge of lawn (1 female); in mailbox (1 female); 

n. d., coastal spruce-mixed hardwood (1 male, 1 female); n. d., saltmarsh (1 female).

Regional Distribution: ME (Collins et al. 1996); ON (Chamberlin and Ivie 1941); QC (Paquin and 

Dupérré 2003).
78

Agelenopsis potteri (Blackwall, 1846)

Taxonomy: Chamberlin and Ivie (1941); Kaston (1981); Roth and Brown (1986); Paquin and 

Dupérré (2003).

Records: M01 (1 male, 20 females); M06 (1 male, 1 female); M21 (3 females). N = 26 adults, 

2 males, 24 females.

Methods: pitfall traps (1 male, 1 female); search (1 male, 20 females); sweep net (3 females).

Months: August (2 females); September (2 males, 16 females); October (6 females).

Habitats: in funnel webs on meadowsweet, old fi eld (2 females); in folded blackberry leaf, old fi eld 

(1 female); in funnel web on blackberry bush, old fi eld (1 female); in funnel web spun between 

goldenrod and blackberry bush, old fi eld (1 female); in vegetable garden (1 female); in web, low grass 

at lawn edge, old fi eld (1 female); in funnel web, base of crabapple tree trunk (1 female); on and 

under bark of dead tree, mixed hardwood-conifer (1 female); sweeping old fi eld (2 females); sweeping 

aster, goldenrod, small alders, old fi eld (1 female); litter, saltmarsh (1 male, 1 female); in mailbox 

(3 females); in web inside bedroom window, house (1 female); in corner, house wall (1 female); in silk 

retreat, split log in woodbin (1 female); inside window, house (1 male); on side of house (2 females); 

in lumber pile (3 females).

Regional Distribution: ME (Procter 1938); NH, MA, QC, NS (Chamberlin and Ivie 1941); CT 

(Kaston 1981).

Agelenopsis utahana (Chamberlin & Ivie, 1933)

Taxonomy: Chamberlin and Ivie (1941); Kaston (1981); Roth and Brown (1986); Paquin and 

Dupérré (2003).

Records: M01 (46 males, 23 females); M04 (2 males, 4 females); M06 (2 males, 1 female); M12 

(1 male, 1 female); M13 (1 female); M18 (1 female); M22 (4 males, 3 females); M29 (1 female); 

M34 (2 females). N = 92 adults; 55 males, 37 females.

Methods: beating cloth (1 female); pitfall traps (40 males, 7 females); search (11 males, 23 females); 

sweep net (3 males, 5 females); n. d. (1 male, 1 female).

Months: July (13 males); July-August (6 males, 2 females); August (28 males, 30 females); AugustSeptember (1 male); September (7 males, 2 females); October (1 female); November (2 females).

Habitats: litter, Betula-Acer stand (30 males, 4 females); litter, bigtooth aspen stand (4 males, 3 

females); litter, saltmarsh (2 males); litter, seashore-backshore (2 males); litter, edge of old fi eld and 

alder swamp (1 male); litter, coastal red spruce stand (1 male); sweeping foliage, freshwater pond 

edge (2 males); sweeping vegetation, old fi eld (1 female); sweeping vegetation, saltmarsh (1 female); 

sweeping vegetation, freshwater marsh (1 male, 1 female); sweeping grasses, sedges, forbs, dry 

streambed (2 females); beating hardwoods (1 female); in webs on aster, old fi eld (2 females); in web, 

rolled aster leaf, old fi eld (1 male); in web on aster, roadside, mixed conifer-hardwood (1 female); 
79

in web, folded blackberry leaves (2 females); in funnel web on sarsparilla sp., coastal spruce-mixed 

hardwood (1 female); in folded sarsparilla leaf, roadside, mixed conifer (1 female); in funnel webs, 

understory spruce-mixed hardwood (2 females); in web, folded alder leaf (1 female); in nests, rolled 

alder leaves (2 females); in folded wild cherry leaf (1 female); in web, wild rose (Rosa sp.) leaf (1 

female); in webs, folded (rolled) leaves of bush- honeysuckle (2 males); in web, between leaves of 

small shrub (1 male); in web, among woodland foliage (1 female); in grasses, old fi eld edge (1 male); 

under bark of white (paper) birch (2 males); on foliage, green bean plant, garden (1 female); running 

on ground, vegetable garden (1 male, 1 female); in funnel webs on potato plants, garden (2 females); 

in web, overturned canoe (1 female); in garage (1 female); in shed (1 male); in bathroom, house 

(1 male); on bathroom ceiling (1 female); on porch (1 male); on outside wall of house (1 female); 

n. d. (1 male, 1 female).

Regional Distribution: ME, NH, MA, RI, NY (Chamberlin and Ivie 1941); CT (Kaston 1981); 

QC (Bélanger and Hutchinson 1992).

Tegenaria domestica (Clerck, 1757)

Taxonomy: Roth (1968); Kaston (1981); Paquin and Dupérré (2003).

Records: M01 (12 males, 3 females); M52 (2 females). N = 17 adults; 12 males, 5 females.

Method: search (12 males, 5 females).

Months: May (2 males); June (4 males, 3 females); July (1 male); August (1 male); September 

(1 male, 2 females); October (3 males).

Habitats: in bathtub, house (8 males, 1 female); in bathroom basin, house (1 male); in funnel webs, 

house cellars (1 male, 2 females); in stored storm door, house basement (1 male); in garage, house 

(1 male, 1 female); in folded blackberry leaf, old fi eld (1 female).

Regional Distribution: ME (Procter 1946, as Tegenaria derhami (Scopoli, 1763)); CT (Kaston 

1981); NY (Crosby and Bishop 1928); QC (Bélanger and Hutchinson 1992).
80

FAMILY HAHNIIDAE

The hahniids spin small delicate sheet webs near the ground; the webs are diffi cult to see unless 

covered by dew or mist from fog. The webs of some species remain to be discovered (Opell and 

Beatty 1976). Some authors indicate that hahniid webs lack a retreat; however, Dondale et al. (1997) 

noted that, in some instances, a funnel retreat may be present.

The hahniids are three-clawed spiders with leg spines and trichobothria similar to the Agelenidae; 

in fact, hahniids were formerly regarded as a subfamily of the Agelenidae (Comstock 1948, Kaston 

1981). Most hahniids are easily recognized and distinguished from other spiders by their transverse 

arrangement of the spinnerets. The three pairs of spinnerets are arranged with their basal segments 

nearly in a straight line. The anterior eyes are arranged in two slightly procured rows; sometimes the 

anterior median eyes are minute, e.g., species of Hahnia. Other family characters, notes on natural 

history, and a key to genera in North America north of Mexico are provided by Bennett (2005b).

Opell and Beatty (1976) revised the Nearctic Hahniidae, including 3 genera and 28 species. 

Since their revision, additional genera and species have been added to the Hahniidae; see Bennett 

(2005b) and Platnick (2007). Descriptions of Cryphoeca montana Emerton 1909, a New England to 

midwestern species, are given by Levi and Field (1954), Kaston (1981), and Roth (1994). Currently, 

the hahniid fauna in Maine consists of four genera and fi ve species; three genera and four species have 

been found in Milbridge. 

Antistea brunnea (Emerton, 1909)

Taxonomy: Gertsch (1934); Opell and Beatty (1976); Paquin and Dupérré (2003).

Records: M12 (23 males, 4 females). N = 27 adults; 23 males, 4 females.

Method: pitfall traps (23 males, 4 females).

Months: June (1 female); June-July (1 female); July (2 females); August (1 male); August-September 

(13 males); September (9 males).

Habitat: freshwater marsh edge (23 males, 4 females).

Regional Distribution: ME (Emerton 1915, as Hahnia burnnea Emerton); CT, NY, ON, QC, NS 

(Opell and Beatty 1976).

Cryphoeca montana Emerton, 1909

Taxonomy: Levi and Field (1954); Kaston (1981); Roth (1994); Paquin and Dupérré (2003).

Records: M01 (3 males, 1 female); M04 (70 males); M07 (1 male); M27 (1 male). N = 76 adults; 75 

males, 1 female.

Methods: beating cloth (1 female); pitfall traps (75 males).

Months: May-June (48 males); June (27 males); October (1 female).
81

Habitats: beating alders, old fi eld edge (1 female); litter, deciduous-coniferous woodland (3 males); 

litter, coastal red spruce stand (70 males); litter, Kalmia-Vaccinium heath (1 male); litter, red maplewhite (paper) birch stand (1 male).

Regional Distribution: ME, NH, VT, MA, NF (Emerton 1914); NY (Crosby and Bishop 1928); 

CT (Kaston 1981); QC (Bélanger and Hutchinson 1992).

Hahnia cinerea Emerton, 1890

Taxonomy: Gertsch (1934); Opell and Beatty (1976); Kaston (1981); Paquin and Dupérré (2003).

Records: M24 (3 males, 1 female); M25 (5 males); M26 (1 male); M27 (1 male). N = 11 adults; 10 

males, 1 female.

Method: pitfall traps (10 males, 1 female).

Months: May (7 males); May-June (2 males, 1 female); June (1 male).

Habitats: forest-fl oor litter, red maple sapling stand (3 males, 1 female); forest-fl oor litter, white 

(paper) birch stand (5 males); marsh litter, saltmarsh edge (1 male); forest-fl oor litter, red maple-white 

birch stand (1 male).

Regional Distribution: ME (Procter 1933); NH, MA, CT, NY, ON, QC, NS (Opell and Beatty 

1976).

Neoantistea agilis (Keyserling, 1887)

Taxonomy: Gertsch (1934); Opell and Beatty (1976); Kaston (1981); Paquin and Dupérré (2003).

Records: M01 (2 males); M06 (4 males, 9 females); M07 (6 females); M11 (1 male, 1 female); M22 

(2 females); M25 (1 male, 3 females); M26 (2 males). N = 31 adults; 10 males, 21 females.

Methods: pitfall traps (10 males, 20 females); sifted litter-hand sorted (1 female).

Months: May (2 males, 4 females); May-June (2 females); June (1 male, 4 females); June-July 

(1 female); July (1 male, 7 females); August (4 males, 1 female); August-September (1 male); 

September (1 male, 2 females).

Habitats: litter, old fi eld-alder swamp edge (2 males); litter, saltmarsh (4 males, 9 females); litter, 

Kalmia-Vaccinium heath (5 females); sifted heath litter, Kalmia-Vaccinium heath (1 female); ground, 

gravel pit, conifer-mixed hardwood opening (1 male, 1 female); forest-fl oor litter, bigtooth aspen 

stand (2 females); forest-fl oor litter, white (paper) birch stand (1 male, 3 females); litter, saltmarsh 

edge (2 males).

Regional Distribution: ME (Bryant 1908, as Hahnia agilis Keyserling); NH, VT, MA, CT, RI, NY, 

ON, QC, NS (Opell and Beatty 1976). 
82

Neoantistea magna (Keyserling 1887)

Taxonomy: Gertsch (1934); Opell and Beatty (1976); Kaston (1981); Paquin and Dupérré (2003).

Records: M01 (227 males, 8 females); M04 (13 males, 22 females); M06 (6 males); M12 (1 female); 

M24 (1 female); M27 (1 male, 4 females). N = 283 adults; 247 males, 36 females.

Method: pitfall traps (247 males, 36 females).

Months: May (1 female); May-June (1 female); June (12 females); June-July (4 females); July (46 

males, 12 females); July-August (50 males); August (116 males, 6 females); August-September (22 

males); September (13 males).

Habitats: forest-fl oor litter, Betula-Acer stand (226 males, 7 females); litter, deciduous-coniferous 

woodland (1 male, 1 female); forest-fl oor litter, coastal red spruce stand (13 males, 22 females); litter, 

saltmarsh (6 males); litter, freshwater marsh edge (1 female); forest-fl oor litter, red maple sapling 

stand (1 female); forest-fl oor litter, red maple-white birch stand (1 male, 4 females).

Regional Distribution: ME (Bryant 1908, as Hahnia radula Emerton); NH, VT, MA, CT, NY, ON, 

QC, NS, NF (Opell and Beatty 1976).
83

FAMILY DICTYNIDAE

Most members of this family are small spiders that spin aerial webs in forbs, shrubs, and trees; a few 

live in leaf debris on the ground. They are among the hackle-band weavers or cribellate spiders that 

spin dense meshes of silk. A series of curved bristles on metatarsus IV, called the calamistrum, is used 

to card silk from a special spinning plate, the cribellum, which lies anterior to the spinnerets. Species 

of Cicurina lack a cribellum and calamistrum.

Webs of some dictynids are sheet-like and can be found on walls of buildings, fences, and tree 

bark; others spin tangle-like webs near the apices of grasses, forbs, and dead twigs. Female dictynids 

generally produce more than one egg sac, which are hung within the confi nes of their webs. After 

mating, male dictynids are often seen cohabiting with the females in their web.

Dictynid spiders generally have eight eyes arranged in two rows, or six eyes in two triads, with the 

anterior median eyes missing. The number of eyes may be further reduced or completely absent in 

cave-dwelling Cicurina, a genus formerly assigned to the Agelenidae. Unlike most aerial-spinning 

dictynids, species of Cicurina generally live under rocks or in litter; little is known about their webspinning habits. Additional information on dictynid character descriptors, natural history, and key to 

the genera of North America can be found in Bennett (2005a).

Earlier, Chamberlin and Gertsch (1958) provided keys, illustrations, and descriptions for most 

species of Dictynidae found in North America north of Mexico. However, numerous name changes 

have taken place since this earlier revision; see Bennett (2005a) for a review of taxonomic history. 

For example, the subgenus Emblyna has been elevated to full generic level, with the resultant shift in 

numerous species. Platnick (2007) should be consulted for the latest placement of genera and species.

Pierre Paquin and James C. Cokendolpher are currently revising the genus Cicurina, including species 

found in Maine. Until this revision appears, earlier descriptions and illustrations of Cicurina species 

can be found in Chamberlin and Ivie (1940). Currently, the dictynid fauna of Maine comprises 7 

genera and 22 species; 5 genera and 13 species have been found in Milbridge.

Argenna obesa Emerton, 1911

Taxonomy: Chamberlin and Gertsch (1958); Kaston (1981); Paquin and Dupérré (2003).

Records: M01 (1 male, 1 female); M56 (1 female). N = 3 adults; 1 male, 2 females.

Method: litter condenser-Berlese funnel (1 male, 2 females).

Months: June (1 male, 1 female); September (1 female).

Habitats: sifted rockweed debris, seashore (1 male, 1 female); sifted litter (grasses, forbs, tidal debris), 

salt meadow (1 female).

Regional Distribution: ME (Emerton 1914); NH, MA, CT, NY, ON, NF (Chamberlin and 

Gertsch 1958); QC (Bélanger and Hutchinson 1992).
84

Cicurina arcuata Keyserling, 1887

Taxonomy: Chamberlin and Ivie (1940); Paquin and Dupérré (2003).

Records: M01 (2 males).

Method: pitfall traps (2 males).

Months: May-June (1 male); June (1 male).

Habitat: litter, deciduous-coniferous woodland (2 males).

Regional Distribution: ME (Procter 1933); MA, NY, QC (Chamberlin and Ivie 1940).

Cicurina brevis (Emerton, 1890)

Taxonomy: Chamberlin and Ivie (1940); Kaston (1981); Paquin and Dupérré (2003).

Records: M01 (1 male, 1 female); M04 (3 females); M07 (1 female); M22 (1 male, 1 female); M24 

(3 females); M25 (1 female); M27 (1 female). N = 13 adults; 2 males, 11 females.

Methods: pitfall traps (1 male, 11 females); search (1 male).

Months: March (1 male); May (1 female); May-June (2 females); June (2 females); July (2 females); 

July-August (2 females); August (2 females); September (1 male).

Habitats: litter, coastal red spruce stand (3 females); litter, red maple sapling stand (3 females); litter, 

bigtooth aspen stand (1 male, 1 female); litter, white (paper) birch stand (1 female); litter, red maplewhite (paper) birch stand (1 female); litter, Kalmia-Vaccinium heath (1 female); on ground, BetulaAcer stand (1 female); in container of water (alive) (1 male).

Regional Distribution: ME (Procter 1933); NH, MA, CT, NY (Chamberlin and Ivie 1940); QC 

(Bélanger and Hutchinson 1992).

Cicurina pallida Keyserling, 1887

Taxonomy: Chamberlin and Ivie (1940); Kaston (1981); Paquin and Dupérré (2003).

Records: M01 (1 male, 1 female); M04 (1 male, 1 female); M22 (1 female). N = 5 adults; 2 males, 

3 females.

Methods: pitfall traps (1 male, 3 females); search (1 male).

Months: May (1 male); May-June (1 male); June (1 female); August-September (1 female); October 

(1 female).

Habitats: litter, coastal red spruce stand (1 male, 1 female); litter, bigtooth aspen stand (1 female); 

litter, edge of old fi eld and alder swamp (1 female); in wood pile (1 male).
85

Regional Distribution: ME (Procter 1946); NH, MA, NY (Chamberlin and Ivie 1940); CT (Kaston 

1981); QC (Bélanger and Hutchinson 1992).

Cicurina placida Banks, 1892

Taxonomy: Chamberlin and Ivie (1940).

Record: M01 (1 male). N = 1 adult male.

Method: pitfall trap (1 male).

Month: June (1 male).

Habitat: litter, deciduous-coniferous woodland (1 male).

Regional Distribution: ME (Jennings et al. 1988); NH, NY (Chamberlin and Ivie 1940); CT 

(Kaston 1981).

Dictyna brevitarsa Emerton, 1915

Taxonomy: Chamberlin and Gertsch (1958, as Dictyna brevitarsus Emerton); Kaston (1981, as 

Dictyna brevitarsus Emerton); Paquin and Dupérré (2003, as Dictyna brevitarsus Emerton).

Records: M01 (1 male, 2 females); M04 (1 female); M06 (2 males, 2 females); M10 (1 male); M15 

(7 males). N = 16 adults; 11 males, 5 females.

Method: beating cloth (11 males, 5 females).

Months: May (4 males); June (7 males, 1 female); July (2 females); August (1 female); October (1 

female).

Habitats: beating red spruce foliage, near saltmarsh (7 males); beating red spruce foliage, old fi eld 

edge (1 male, 1 female); beating red spruce foliage, seashore-backshore (1 female); beating red spruce 

foliage, saplings, conifer-mixed hardwood (1 male); beating white spruce foliage, saltmarsh edge 

(2 males, 2 females); beating spruces, old fi eld edge (1 female).

Regional Distribution: ME, NH, VT, MA, CT, NY, ON, QC, NS, NF (Chamberlin and Gertsch 

1958, as Dictyna brevitarsus Emerton).

Dictyna coloradensis Chamberlin, 1919

Taxonomy: Chamberlin and Gertsch (1958); Kaston (1981); Paquin and Dupérré (2003).

Records: M01 (1 male, 15 females); M06 (1 female); M35 (2 females); M51 (1 female). N = 20 

adults; 1 male, 19 females.

Methods: search (1 male, 15 females); sweep net (3 females); n. d. (1 female).
86

Months: June (1 female); July (1 male, 5 females); August (8 females); September (4 females); 

October (1 female).

Habitats: in webs on dry seedheads of meadowsweet (1 male, 2 females); in webs on meadowsweet, 

old fi eld (2 females); in nest, top of Spiraea twig (1 female); in web, dried seedhead of Hieracium sp. 

(1 female); in webs, grass seedheads, old fi eld and n. d. (2 females); in silken retreat, folded Vaccinium

leaf, old fi eld (1 female); in web, fl at-topped goldenrod, old fi eld (1 female); in web, dead Amelanchier 

seedling (1 female); in web, apex of dry forb, old fi eld (1 female); in shrubs, old fi eld (1 female); in 

webs, dry rush (Juncus sp.) seedheads, brackish marsh (2 females); sweeping vegetation of old fi eld 

(2 females); sweeping meadowsweet, mixed conifer opening (1 female); n. d., saltmarsh (1 female).

Regional Distribution: ME, NH, VT, MA, CT, RI, NY, ON, QC (Chamberlin and Gertsch 1958).

Dictyna minuta Emerton, 1888

Taxonomy: Chamberlin and Gertsch (1958); Kaston (1981); Paquin and Dupérré (2003).

Record: M06 (1 female). N = 1 adult female.

Method: sweep net (1 female).

Month: August (1 female).

Habitat: sweeping saltmarsh vegetation (1 female).

Regional Distribution: ME (Jennings et al.1990); NH, MA, CT, RI, NY (Chamberlin and Gertsch 

1958); QC (Bélanger and Hutchinson 1992).

**Emblyna manitoba (Ivie, 1947)

Taxonomy: Chamberlin and Gertsch (1958); Paquin and Dupérré (2003).

Records: M06 (4 females). N = 4 adult females.

Method: sweep net (4 females).

Month: June (4 females).

Habitat: sweeping saltmarsh vegetation (4 females).

Regional Distribution: ME (this study); ON (Chamberlin and Gertsch 1958); QC (Bélanger and 

Hutchinson 1992).

Emblyna maxima (Banks, 1892)

Taxonomy: Chamberlin and Gertsch (1958); Kaston (1981); Paquin and Dupérré (2003).

Records: M01 (4 females); M14 (1 male). N = 5 adults; 1 male, 4 females.
87

Methods: search (1 male); sweep net (4 females).

Months: June (2 females); July (1 male, 1 female); August (1 female).

Habitats: sweeping grasses and forbs, mixed hardwood-conifer (1 male); sweeping low vegetation, 

deciduous-coniferous woodland (1 female); sweeping Vaccinium and small shrubs, old fi eld (1 

female); sweeping low shrubs, Betula-Acer understory (1 female); in web on alder twig (1 female).

Regional Distribution: ME (Procter 1946, as Dictyna maxima Banks); NH, VT, MA, CT, NY, ON, 

QC (Chamberlin and Gertsch 1958, as Dictyna maxima Banks).

Emblyna phylax (Gertsch & Ivie, 1936)

Taxonomy: Chamberlin and Gertsch (1958); Paquin and Dupérré (2003).

Records: M01 (12 males, 2 females); M05 (1 male); M06 (5 males, 2 females); M10 (1 male); M13 

(1 female); M15 (5 males); M31 (1 female); M32 (2 females); M51 (2 females). N = 34 adults; 24 

males, 10 females.

Methods: beating cloth (22 males, 10 females); search (1 male); sweep net (1 male).

Months: May (17 males); June (7 males, 5 females); July (4 females); August (1 female).

Habitats: beating red spruce foliage, old fi eld edge (11 males, 2 females); beating red spruce foliage, 

mixed hardwood-conifer (2 females); beating red spruce foliage, near saltmarsh (5 males); beating red 

spruce foliage, mixed conifer (2 females); beating red spruce foliage, white pine-red spruce (1 female); 

beating white spruce foliage, saltmarsh edge (5 males, 2 females); beating eastern larch sapling foliage, 

mixed conifer-hardwood (1 male); beating balsam fi r foliage, mixed hardwood-conifer (1 female); 

sweeping wet meadow, old fi eld-brackish marsh edge (1 male); in box, carpenter’s debris (1 male).

Regional Distribution: ME (Procter 1946, as Dictyna phylax Gertsch & Ivie); NH, NY, ON, NF 

(Chamberlin and Gertsch 1958, as Dictyna phylax Gertsch & Ivie); QC (Bélanger and Hutchinson 

1992).

Emblyna sublata (Hentz, 1850)

Taxonomy: Chamberlin and Gertsch (1958); Kaston (1981); Paquin and Dupérré (2003).

Record: M35 (1 female). N = 1 adult female.

Method: sweep net (1 female).

Month: June (1 female).

Habitat: sweeping grasses and rushes, brackish marsh (1 female).
88

Regional Distribution: ME (Procter 1933, as Dictyna sublata (Hentz)); NH, VT, MA, CT, NY, ON 

(Chamberlin and Gertsch 1958, as Dictyna sublata (Hentz)); QC (Bélanger and Hutchinson 1992, as 

Dictyna sublata (Hentz)).

Lathys pallida (Marx, 1891)

Taxonomy: Chamberlin and Gertsch (1958); Kaston (1981); Paquin and Dupérré (2003).

Records: M24 (1 male); M25 (5 males, 5 females); M27 (5 males, 1 female); M53 (3 females). N = 

20 adults; 11 males, 9 females.

Methods: litter condenser-Berlese funnel (3 females); pitfall traps (11 males, 6 females).

Months: May (3 males, 4 females); May-June (5 males); June (1 male, 2 females); June-July (1 male); 

July (1 male, 3 females). 

Habitats: litter, white (paper) birch stand (5 males, 5 females); litter, red maple-white birch stand (5 

males, 1 female); litter, red maple sapling stand (1 male); sifted aspen-maple litter, mixed hardwoods 

(3 females).

Regional Distribution: ME (Procter 1946, as Scotolathys pallida (Marx); NH, VT, MA, CT, NY, 

ON (Chamberlin and Gertsch 1958); QC (Bélanger and Hutchinson 1992).
89

FAMILY AMAUROBIIDAE

The amaurobiids common to New England and the northern region are cribellate, or hackle-band 

weavers, that spin dense mesh webs on and under loose bark of trees and fallen logs. Rock fi ssures, 

crevices, woodpiles, caves, and cellars are also favored habitats of amaurobiids. In Maine, these spiders 

are frequently brought indoors on fi rewood; however, they are non-aggressive and pose no known 

medical problems.

Amaurobiids have three tarsal claws, eyes arranged in two rows of four each, and robust chelicerae. 

Like most dictynids, our northern amaurobiids have a cribellum and calamistrum; the cribellum is 

divided into two halves. Ubick (2005a) describes other characters of this family and provides notes on 

natural history. He also gives a key to the genera of Amaurobiidae found in North America north of 

Mexico.

Two genera, Coras and Wadotes, formerly assigned to the Agelenidae, have been shifted to 

the Amaurobiidae, and one amaurobiid subfamily has been elevated to full family status, the 

Titanoecidae. With these recent taxonomic changes, the Maine amaurobiid fauna is now represented 

by 5 genera and 10 species. The Milbridge amaurobiid fauna consists of fi ve genera, each represented 

by only one species.

Amaurobius borealis Emerton, 1909

Taxonomy: Leech (1972); Kaston (1981); Paquin and Dupérré (2003).

Records: M07 (1 male); M24 (2 males); M25 (19 males, 3 females); M27 (13 males, 1 female). N = 

39 adults; 35 males, 4 females.

Method: pitfall traps (35 males, 4 females).

Months: May (22 males, 3 females); May-June (10 males); June (3 males); June-July (1 female).

Habitats: litter, Kalmia-Vaccinium heath (1 male); forest-fl oor litter, red maple sapling stand 

(2 males); forest-fl oor litter, white (paper) birch stand (19 males, 3 females); forest-fl oor litter, red 

maple-white birch stand (13 males, 1 female).

Regional Distribution: ME (Emerton 1914, as Walmus borealis (Emerton)); MA, NH, VT, ON, 

QC, NB, NF, LB (Leech 1972).

Callobius bennetti (Blackwall, 1846)

Taxonomy: Leech (1972); Kaston (1981); Paquin and Dupérré (2003).

Records: M01 (5 males, 8 females); M04 (15 males, 14 females); M07 (2 males); M10 (1 male); 

M13 (1 male); M22 (1 female); M23 (9 females); M24 (1 female); M25 (7 males, 5 females); M27 

(11 males, 3 females). N = 83 adults; 42 males, 41 females.

Methods: beating cloth (1 female); pitfall traps (35 males, 20 females); search (7 males, 19 females); 

sweep net (1 female).
90

Months: March (1 female); May (20 males); May-June (9 males, 2 females); June (12 males, 10 

females); June-July (7 females); July (1 male, 14 females); August (5 females); September (1 female); 

October (1 female).

Habitats: beating foliage, hardwoods (1 female); forest-fl oor litter, Betula-Acer stand (1 female); 

forest-fl oor litter, coastal red spruce stand (15 males, 9 females); litter, Kalmia-Vaccinium heath 

(2 males); forest-fl oor litter, bigtooth aspen stand (1 female); forest-fl oor litter, red maple sapling 

stand (1 female); forest-fl oor litter, white (paper) birch stand (7 males, 5 females); forest-fl oor litter, 

red maple-white birch stand (11 males, 3 females); sweeping vegetation, old fi eld (1 female); in 

birch, coastal spruce-mixed hardwood (1 female); under loose bark, white (paper) birch, mixed 

conifer-hardwood (2 females); under bark, dead white (paper) birch, coastal spruce-mixed hardwood 

(1 female); under bark, dead tree (n. d.), coastal red spruce stand (1 female); under rocks on ledge, 

mixed conifer-hardwood (1 male); in web, bridge guardrail, prey of Achaearanea tabulata female, 

roadside-riparian (1 male); under rocks, mixed conifer, island (7 females); under bark, white (paper) 

birch, island (1 female); under loose bark, dead birch, island (1 female); under bark, dead tree, mixed 

hardwood-conifer (1 female); under log, deciduous-coniferous woodland (1 female); in bathtub, 

house (3 males); in bathroom basin, house (1 male, 1 female); in kitchen sink, house (1 female); on 

rug, house bedroom (1 female); on living room wall, house (1 male).

Regional Distribution: ME (Bishop 1923); NH, VT, CT, NY, VT, NB, NF, NS, ON, QC (Leech 

1972).

Coras montanus (Emerton, 1890)

Taxonomy: Muma (1946); Kaston (1981); Roth (1994); Paquin and Dupérré (2003).

Records: M01 (2 males, 1 female); M04 (2 males); M22 (1 male); M24 (1 male); M27 (1 male); 

M34 (1 female). N = 9 adults; 7 males, 2 females.

Methods: pitfall traps (3 males); search (4 males, 2 females).

Months: April (1 male); May-June (1 male); June (2 males, 1 female); July (1 male); August (1 male, 

1 female); September (1 male).

Habitats: forest-fl oor litter, bigtooth aspen stand (1 male); forest-fl oor litter, red maple sapling stand 

(1 male); forest-fl oor litter, red maple-white birch stand (1 male); under loose bark, white (paper) 

birch (1 male); under curled bark, white (paper) birch, mixed conifer-hardwood (1 female); under 

bark, dead spruce, seashore (1 male); in kitchen sink (dead), house (1 male); in Jacuzzi fi lter (dead) (1 

male); in tarpaulin, screen house (1 female).

Regional Distribution: ME (Procter 1938); NH, CT, NY, ON, QC, NF, NS (Muma 1946).

Cybaeopsis euopla (Bishop & Crosby, 1935)

Taxonomy: Bishop and Crosby (1935, as Callioplus euoplus, n. sp.); Leech (1972, as Callioplus 

euoplus Bishop and Crosby); Paquin and Dupérré (2003). This species was formerly assigned to the 

Dictynidae by Bishop and Crosby (1935).
91

Records: M12 (2 males, 1 female). N = 3 adults; 2 males, 1 female.

Method: pitfall traps (2 males, 1 female).

Months: June (2 males); September (1 female).

Habitat: litter, freshwater marsh edge (2 males, 1 female).

Regional Distribution: ME (Bishop and Crosby 1935); ON, QC, NF, NS (Leech 1972).

Wadotes calcaratus (Keyserling, 1887)

Taxonomy: Muma (1947); Kaston (1981); Bennett (1987); Paquin and Dupérré (2003). Muma 

(1947), Kaston (1981), and others formerly assigned this species to the Agelenidae.

Records: M04 (9 males); M24 (1 male, 1 female); M27 (2 males). N = 13 adults; 12 males, 1 female.

Method: pitfall traps (12 males, 1 female).

Months: May-June (4 males, 1 female); June (4 males); August (4 males).

Habitats: forest-fl oor litter, coastal red spruce stand (9 males); forest-fl oor litter, red maple sapling 

stand (1 male, 1 female); forest-fl oor litter, red maple-white (paper) birch stand (2 males).

Regional Distribution: ME (Chamberlin 1925); NH, VT, MA, CT, NY, ON, QC, NB, NS (Roth 

and Brown 1986; Bennett 1987).
92

FAMILY TITANOECIDAE

The titanoecids were formerly included as a subfamily, the Titanoecinae, of the family Amaurobiidae. 

They share many characters in common with amaurobiids; however, the leg trichobothria are short 

and thick and do not increase in length distally. Their habitats are similar to those of amaurobiids, 

but chiefl y under rocks, logs, and bark on the ground. Cutler (2005) provides additional character 

descriptors and brief notes on the natural history of this small family. A key to the North American 

species is found in Leech (1972), who treated the Titanoecidae as a subfamily of the Amaurobiidae.

Thus far, the Maine titanoecid fauna is represented by only one species, Titanoeca americana Emerton 

1888. However, one additional species, Titanoeca brunnea Emerton 1888, possibly occurs in the 

State; it has been taken in Connecticut, Massachusetts, New Hampshire, New York, and Ontario. 

Descriptions and illustrations of both species can be found in Leech (1972).

Titanoeca americana Emerton, 1888

Taxonomy: Lehtinen (1967); Leech (1972); Paquin and Dupérré (2003).

Record: M04 (1 female). N = 1 adult female.

Method: search (1 female).

Month: August (1 female).

Habitat: under rock, upper shore ledges, seashore (1 female).

Regional Distribution: ME (Procter 1933, as Amaurobius americanus (Emerton)); MA, CT, NH, 

NY, ON (Leech 1972); QC (Bélanger and Hutchinson 1992).

Note: The only other known locality for this species in Maine is Mount Desert Island, Hancock 

County. On Mount Desert Island, Procter (1933) noted that this species was taken under stones in 

a very hot, dry area (i.e., “The Hop”); Leech (1972) recorded it from Champlain Mountain at an 

altitude of 330 m.
93

ANNOTATED FAUNAL LIST: 

HUNTER FAMILIES

FAMILY LYCOSIDAE

The lycosids or wolf spiders are among the most commonly observed spiders in Maine. They are 

usually seen running through grass of old fi elds, or among dead leaves on the forest fl oor, or over 

sandy and stony places along streams, lakes, and cobble beaches. Most are cursorial hunters that 

detect and capture prey by “sit and wait” tactics. With few exceptions, snares and retreats are not 

made; however, species of Geolycosa line their tubular tunnels with silk.

In this family the eyes are unequal in size and appear to be arranged in three rows. The anterior eyes 

are the smallest and are aligned in a row of four above the clypeus. The posterior row is strongly 

recurved, thus forming two rows of two eyes each. The posterior median eyes (i.e., the second row) 

are the largest by far. Many lycosids are active at night as well as during daylight hours. The females 

characteristically carry their egg sacs attached to the spinnerets. Dondale (2005a) provides additional 

information on family-character descriptors, natural and taxonomic histories, and a key to the lycosid 

genera of North America north of Mexico.

Most of the lycosid species found in the northern region can be identifi ed using the identifi cation 

keys, descriptions, and illustrations provided by Dondale and Redner 1990. Earlier revisionary works 

by these same authors are also helpful; see Dondale and Redner (1990) for citations. The lycosid 

fauna in Maine is represented by 10 genera and 48 species; 8 genera and 25 species, including a new 

species of Pirata, make up the lycosid fauna in Milbridge.

Alopecosa aculeata (Clerck, 1757)

Taxonomy: Dondale and Redner (1979, 1990); Kaston (1981); Paquin and Dupérré (2003).

Records: M01 (8 males, 2 females); M04 (1 female); M06 (1 female); M07 (2 males); M14 (1 male); 

M22 (1 male, 6 females); M23 (1 female); M24 (3 males); M25 (11 males, 1 female); M26 (5 males); 

M27 (12 males, 1 female); M58 (1 male, 1 female). N = 58 adults; 44 males, 14 females.

Methods: pitfall traps (39 males, 9 females); search (5 males, 5 females).

Months: May (7 males, 2 females); May-June (13 males); June (23 males, 2 females); July (1 male, 4 

females); July-August (2 females); August (3 females); September (1 female).

Habitats: litter, seashore-backshore (4 males); litter, deciduous-coniferous woodland (1 male); litter, 

saltmarsh (1 female); litter, Kalmia-Vaccinium heath (2 males); litter, bigtooth aspen stand (1 male, 

6 females); litter, red maple sapling stand (3 males); litter, white (paper) birch stand (11 males, 1 

female); litter, saltmarsh edge (5 males); litter, red maple-white birch stand (12 males, 1 female); 

in grass near shore, seashore (1 male); on ground, coastal red spruce-mixed hardwoods (1 female); 

on ground among leaves and grass, mixed hardwood-conifer (1 male); on ledge under fl aked stone, 

mixed conifer, island (1 female); on ground among grass and matted straw, seashore-brackish marsh 

(1 male, 1 female); running in vegetable garden (1 female); running in compost pile (1 male); on 

outside wall of house (1 male); on bathroom fl oor, house (1 female).
94

Regional Distribution: ME (Emerton 1914, as Lycosa beanii Emerton); NH, CT, NY, QC, NB, NS, 

NF (Dondale and Redner 1979).

Arctosa emertoni Gertsch, 1934

Taxonomy: Dondale and Redner (1983, 1990); Kaston (1981); Paquin and Dupérré (2003).

Records: M05 (1 female); M06 (11 males, 9 females); M23 (1 female); M26 (32 males, 16 females). 

N = 70 adults; 43 males, 27 females.

Methods: pitfall traps (43 males, 25 females); search (2 females).

Months: May (3 males, 4 females); May-June (11 males, 4 females); June (20 males, 8 females); 

June-July (1 male); July (7 males, 6 females); July-August (3 females); August (1 male, 1 female); 

September (1 female).

Habitats: litter, saltmarsh (11 males, 9 females); litter, saltmarsh edge (32 males, 16 females); 

burrowed under board, old fi eld-brackish marsh margin (1 female); on sand-rock beach below wrack 

line, seashore, island (1 female).

Regional Distribution: ME, NH, VT, MA, NY, QC, NB, NS (Dondale and Redner 1983); CT 

(Kaston 1981); ON (Dondale and Redner 1990). 

Arctosa rubicunda (Keyserling, 1877)

Taxonomy: Dondale and Redner (1983, 1990); Kaston (1981); Paquin and Dupérré (2003).

Records: M06 (10 males, 8 females). N = 18 adults; 10 males, 8 females.

Method: pitfall traps (10 males, 8 females).

Months: May (4 males), June (3 males); July (3 males, 6 females); August (2 females).

Habitat: litter, saltmarsh (10 males, 8 females).

Regional Distribution: ME (Bryant 1908, as Lycosa rubicunda (Keyserling) Montgomery); NY, ON, 

QC, NB, NS (Dondale and Redner 1983); CT (Kaston 1981); MA (Dondale and Redner 1990).

Hogna frondicola (Emerton, 1885)

Taxonomy: Dondale and Redner (1990); Kaston (1981, as Lycosa frondicola (Emerton)); Paquin and 

Dupérré (2003).

Records: M01 (4 males, 16 females); M05 (1 female); M07 (2 males); M10 (1 female); M14 (1 

female); M15 (1 female); M22 (1 male, 5 females); M25 (3 males); M26 (1 male); M27 (1 male). N 

= 37 adults; 12 males, 25 females.

Methods: pitfall traps (9 males, 6 females); search (3 males, 19 females).
95

Months: April (1 male); May (9 males, 8 females); July (11 females); August (3 females); AugustSeptember (1 female); September (2 males, 2 females).

Habitats: litter, edge of old fi eld and alder swamp (1 male, 1 female); litter, Kalmia-Vaccinium heath 

(2 males); litter, bigtooth aspen stand (1 male, 5 females); litter, white (paper) birch stand (3 males); 

litter, saltmarsh edge (1 male); litter, red maple-white (paper) birch stand (1 male); in soil, vegetable 

garden (1 female); in soil under mulch, vegetable garden (1 female); in vegetable garden (2 females); 

running over soil, vegetable garden (1 female); running through grass, lawn (1 male, 1 female); 

running in or through grass (1 male, 2 females); running at edge of old fi eld (1 female); old fi eld (1 

female); running across forest path, mixed conifer-hardwood (1 female); running across stone terrace 

(1 female); on ground in garden (2 females); on ground in grass (1 female); on ground in mowed 

fi eld (1 female); under rock on ledge, mixed conifer-hardwood (1 female); under litter, upper wrack 

line, saltmarsh (1 female); moribund on terrace, house (1 female); on fl oor of sunroom, house (1 

male).

Regional Distribution: ME (Bryant 1908, as Lycosa frondicola Emerton); NH, VT, MA, NY, ON, 

QC, NB, NS, NF (Dondale and Redner 1990); CT (Kaston 1981, as Lycosa frondicola (Emerton)).

Pardosa distincta (Blackwall, 1846)

Taxonomy: Vogel (1964, 2004); Kaston (1981); Dondale and Redner (1990); Paquin and Dupérré 

(2003).

Records: M01 (7 males, 14 females); M05 (1 female); M06 (2 males, 6 females); M07 (38 males, 22 

females); M11 (1 male, 6 females); M26 (2 males, 1 female); M31 (1 female); M50 (1 female); M51 

(1 male). N = 103 adults; 51 males, 52 females.

Methods: pitfall traps (44 males, 37 females); search (6 males, 15 females); sweep net (1 male).

Months: May (2 males, 1 female); May-June (4 males); June (23 males, 13 females); June-July (4 

males, 6 females); July (18 males, 25 females); August (6 females); September (1 female).

Habitats: litter, Amelanchier-Rubus stand (1 male, 1 female); litter, edge of old fi eld and alder swamp 

(1 female); litter, saltmarsh (2 males, 6 females); litter, Kalmia-Vaccinium heath (38 females, 22 

females); ground, gravel pit, conifer-mixed hardwood (1 male, 6 females); litter, saltmarsh edge (2 

males, 1 female); in reindeer lichen (1 female); in old fi eld (1 male); in garden (1 male); on reindeer 

moss, mixed hardwood-conifer (3 females); on ground in mowed fi eld (1 female); on lawn (1 male); 

on ground in conifer clearing (1 female); on ground, cobble beach, seashore (1 female); on woods 

road, mixed conifer opening (1 male); running on ground, vegetable garden (1 female); running 

through straw, vegetable garden (1 male); running in compost pile (1 male); running on reindeer 

lichen, ledges, mixed conifer-hardwoods (1 female); running through grass (1 female); running in 

old fi eld (2 females); running on rock in old fi eld (1 female); among reindeer lichen on ledges, mixed 

hardwood-conifer (2 females); sweeping vegetation, old fi eld (1 male).

Regional Distribution: ME (Bryant 1908, as Pardosa pallida Emerton); NH, VT, ON, QC, NB, NS 

(Dondale and Redner 1990); CT (Kaston 1981).
96

Pardosa fuscula (Thorell, 1875)

Taxonomy: Dondale and Redner (1987, 1990); Paquin and Dupérré (2003); Vogel (2004).

Records: M12 (5 females). N = 5 adult females.

Method: search (5 females).

Month: June (5 females).

Habitat: on ground, dry matted grasses, freshwater marsh (5 females).

Regional Distribution: ME (Bryant 1908, as Pardosa glacialis (Thorell)); VT, NY, QC, NB, NS, NF, 

LB (Dondale and Redner 1987); ON (Dondale and Redner 1990).

Note: According to J. H. Redner (pers. comm.), early Maine records of Pardosa glacialis (Thorell, 

1872) probably refer to Pardosa fuscula (Thorell, 1875).

Pardosa groenlandica (Thorell, 1872)

Taxonomy: Kronestedt (1975); Dondale and Redner (1990); Paquin and Dupérré (2003); Vogel 

(2004).

Records: M01 (4 males, 28 females); M08 (3 females); M23 (1 male); M50 (1 female).

Methods: pitfall trap (1 female); search (9 males, 27 females). N = 37 adults; 9 males, 28 females.

Months: April (5 males); May (2 males, 6 females); June (1 male, 7 females); July (1 male, 12 

females); August (3 females).

Habitats: among rocks on cobble beach, seashore (3 females); among pebbles and dead rockweed, 

seashore (4 males); among pebbles and grasses on shore, seashore (1 female); on pebbly beach in 

grasses and dead rockweed, seashore (5 females); on rocks at shore, seashore (2 females); on beach 

rocks, seashore, island (1 male); on gravel-pebble beach, seashore (1 male); on pebbled beach, 

seashore (1 female); on rocky shore, seashore (1 female); on bay rocks, seashore (1 female); on large 

rock, bay shore, seashore (1 female); on cobble beach under stone, seashore (1 female); on ground, 

intertidal zone, seashore (1 female); on shingle beach along river, riparian (1 male); running on 

grass and pebbles, cobble beach, seashore (2 females); running on cobble beach, seashore (1 female); 

running through grass and pebbles at shore, seashore (1 female); running on beach, seashore (1 

female); running on sea grasses (rockweed) at edge, seashore (1 male); running on rocks at shore, 

seashore (1 male, 1 female); under small rock at shore, seashore (1 female); under rocks along 

shore, seashore (2 females); under rock, cobble beach, seashore (1 female); litter, seashore-backshore 

(1 female).

Regional Distribution: ME (Bryant 1908); ON, QC, NS, NF, LB (Dondale and Redner 1990).
97

Pardosa hyperborea (Thorell, 1872)

Taxonomy: Dondale and Redner (1987, 1990); Paquin and Dupérré (2003); Vogel (2004).

Records: M04 (2 females); M23 (2 females); M25 (1 male); M42 (1 female); M49 (3 females). N = 9 

adults; 1 male, 8 females.

Methods: pitfall trap (1 male); search (7 females); sweep net (1 female).

Months: June (1 male); July (6 females); August (2 females).

Habitats: litter, white (paper) birch stand (1 male); sweeping Vaccinium sp., island (1 female); on 

ground vegetation near rocky shore, seashore (1 female); on reindeer lichen, ledge near seashore 

(1 female); on Sphagnum moss, island (1 female); running on reindeer lichen, ledges, mixed conifer, 

island (1 female); on reindeer moss, coastal red spruce, upper shore, seashore (1 female); on ground, 

upper shore, seashore-backshore (2 females).

Regional Distribution: ME (Bryant 1908, as Pardosa luteola Emerton); NH, VT, NY, QC, NB, NS, 

NF, LB (Dondale and Redner 1987); ON (Dondale and Redner 1990).

Pardosa lapidicina Emerton, 1885

Taxonomy: Barnes (1959); Kaston (1981); Dondale and Redner (1990); Paquin and Dupérré 

(2003); Vogel (2004).

Records: M01 (1 female); M04 (5 males, 3 females); M49 (4 males, 1 female). N = 14 adults; 

9 males, 5 females.

Methods: search (9 males, 5 females).

Months: May (4 males, 3 females); June (1 male); July (4 males, 1 female); August (1 female).

Habitats: on rocks, cobble beach, seashore (4 males, 1 female); on cobblestone beach and ledge, 

seashore (3 males, 1 female); on rocks at shore, seashore (1 male, 1 female); on ledges, seashore 

(1 male); on pebbly beach in grasses and dead rockweed, seashore (1 female); among rocks on shore, 

seashore (1 female).

Regional Distribution: ME (Bryant 1908); CT (Kaston 1981); NY, ON, QC, NB, NS, NF 

(Dondale and Redner 1990).

Pardosa mackenziana (Keyserling, 1877)

Taxonomy: Lowrie and Dondale (1981); Dondale and Redner (1990); Paquin and Dupérré (2003); 

Vogel (2004).

Records: M01 (3 males, 5 females); M04 (1 male, 2 females); M07 (2 males); M22 (2 males); M24 

(2 males, 1 female); M25 (2 males, 2 females); M27 (1 male, 1 female). N = 24 adults; 13 males, 11 

females.
98

Methods: pitfall traps (13 males, 5 females); search (4 females); sweep net (1 female); n. d. (1 female).

Months: May (2 females); May-June (1 male); June (6 males, 6 females); June-July (2 males); July 

(4 males, 1 female); August (2 females).

Habitats: litter, Amelanchier-Rubus stand (1 male, 1 female); litter, deciduous-coniferous woodland 

(2 males); litter, coastal red spruce stand (1 male); litter, Kalmia-Vaccinium heath (2 males); litter, 

bigtooth aspen stand (2 males); litter, red maple sapling stand (2 males, 1 female); litter, white (paper) 

birch stand (2 males, 2 females); litter, red maple-white birch stand (1 male, 1 female); sweeping 

shrubby vegetation, coastal spruce-mixed hardwood (1 female); on ground, old fi eld (1 female); new 

shore, seashore (1 female); on compost pile, garden (1 female); in cellar bulkhead, house (1 female); 

on outside wall of house (1 female).

Regional Distribution: ME (Bryant 1908, as Pardosa uncata (Thorell)); MA, NH, ON, QC, NB, 

NS, NF (Dondale and Redner 1990).

Note: Early records of Pardosa uncata (Thorell, 1877) in Maine (e.g., Bryant 1908, Emerton 1914, 

and others) most likely refer to Pardosa mackenziana (Keyserling, 1877), a structurally similar but 

widely distributed species. According to Lowrie and Dondale (1981), P. uncata is a western species 

found chiefl y in moist coniferous forests of ID, UT, CO, and NM. More recent, defi nitive records of 

P. mackenziana in Maine include Jennings et al. (1988) and Hilburn and Jennings (1988).

Pardosa modica (Blackwall, 1846)

Taxonomy: Kronestedt (1981); Kaston (1981); Dondale and Redner (1990); Paquin and Dupérré 

(2003); Vogel (2004).

Records: M01 (1 male, 2 females); M06 (105 males, 177 females); M08 (2 females); M17 (2 males, 

1 female); M26 (6 males, 26 females); M58 (3 females). N = 326 adults; 114 males, 212 females.

Methods: pitfall traps (112 males, 198 females); search (11 females); sweep net (2 males, 3 females).

Months: April (1 male, 1 female); May (109 males, 78 females); May-June (1 male, 8 females); June 

(2 males, 54 females); June-July (1 male, 1 female); July (62 females); July-August (3 females); August 

(4 females); August-September (1 female).

Habitats: litter, saltmarsh (105 males, 174 females); litter, saltmarsh edge (6 males, 24 females); 

sweeping grasses, saltmarshes (2 males, 2 females); sweeping vegetation, saltmarsh edge (1 female); 

on ground among marsh vegetation and litter, saltmarsh (2 females); on ground, saltmarsh edge 

(1 female); on ground, brackish marsh (1 female); on ground, seashore-brackish marsh (3 females); on 

rocky beach, seashore (1 female); among shore grasses, seashore (2 females); among pebbles and dead 

rockweed (1 male, 1 female).

Regional Distribution: ME (Procter 1933, as Pardosa brunnea Emerton); CT (Kaston 1981); NY 

(Crosby and Bishop 1928); ON, QC, NB, NS (Dondale and Redner 1990).

Note: Later, Procter (1938, 1946) listed this species as Pardosa modica (Blackwall) (brunnea Emerton).
99

Pardosa moesta Banks, 1892

Taxonomy: Dondale and Redner (1987, 1990); Kaston (1981); Paquin and Dupérré (2003); Vogel 

(2004).

Records: M01 (56 males, 68 females); M05 (1 male, 4 females); M06 (12 males, 21 females); M07 

(22 males, 9 females); M08 (1 female); M12 (34 males, 65 females); M14 (3 males, 4 females); M21 

(1 male); M23 (3 females); M24 (3 males); M25 (2 males, 1 female); M26 (82 males, 54 females); 

M28 (1 female); M35 (3 females); M38 (1 female); M47 (1 female); M49 (1 female); M50 (1 

female); M51 (4 females); M55 (2 males, 2 females); M58 (1 male, 7 females). N = 470 adults; 219 

males, 251 females.

Methods: pitfall traps (190 males, 172 females); search (27 males, 67 females); sweep net (2 males, 

11 females); n. d. (1 female).

Months: May (31 males, 25 females); May-June (17 males, 1 female); June (124 males, 52 females); 

June-July (26 males, 13 females); July (18 males, 73 females); July-August (3 males, 13 females); 

August (58 females); August-September (12 females); September (4 females).

Habitats: litter, Betula-Acer stand near seashore (1 female); litter, edge of old fi eld and alder swamp 

(19 females); litter, seashore-backshore (35 males, 12 females); litter, saltmarsh (12 males, 13 females); 

litter, Kalmia-Vaccinium heath (22 males, 9 females); litter, freshwater marsh edge (34 males, 63 

females); litter, red maple sapling stand (3 males); litter, white (paper) birch stand (2 males, 1 female); 

litter, saltmarsh edge (82 males, 54 females); sweeping Vaccinium sp. and sweet gale, old fi eld (2 

females); sweeping vegetation, old fi eld (2 females); sweeping wet meadow along river, old fi eldbrackish marsh (1 male); sweeping shrubs and grasses, freshwater marsh (1 female); sweeping grasses 

on river shore, riparian (1 female); sweeping grassy area near cabin, mixed conifer opening, island (2 

females); sweeping roadside disturbed area, mixed hardwood-conifer (1 female); sweeping vegetation 

along freshwater stream, riparian (1 female); sweeping grasses and forbs, edge of playground, 

disturbed area (1 male, 1 female); in old fi eld (1 male); in grasses, old fi eld edge (1 female); in grass 

(1 female); in grass with egg sac (1 female); in garden (1 male, 5 females); in vegetable garden (1 

female); in compost pile (1 male, 2 females); in bathtub, house (1 male); around foundation, house 

(1 male); on ground, grass and matted straw, seashore-brackish marsh (3 females); on ground near 

marsh, seashore-brackish marsh (1 female); on ground among litter, old fi eld (1 female); on ground 

in open fi eld, old fi eld (2 females); on ground, saltmarsh (2 females); on ground among marsh 

vegetation and litter, saltmarsh (4 females); on pebbles along beach, seashore (1 female); among shore 

grasses, seashore (1 female); on ground, dry matted vegetation, freshwater marsh (1 female); along 

forest trail, mixed hardwood-conifer (2 males, 1 female); on ground among leaves and grass, mixed 

hardwood-conifer (1 male, 3 females); on ground, Sphagnum wetland near beach, wetland (1 female); 

on ground among dry grass stems, brackish marsh (3 females); on ground, Sphagnum-cranberry-rush 

bog (1 female); on ground, upper shore, seashore-backshore (1 female); on rock, seashore (1 male); on 

woods road, mixed conifer opening (4 females); on ground near freshwater stream, riparian (1 male); 

on ground among dried marsh vegetation, seashore-brackish marsh (1 male, 3 females); on ground, 

old fi eld edge (1 male); on rock, old fi eld (2 females); on soil in vegetable garden (1 male); on ground 

in garden (3 females); on ground and boards in garden (1 female); on garage fl oor, house (1 male); 

on side of house (1 male); on outside shingled wall, house (2 males); on outside wall, shed (1 male); 
100

on exterior vinyl siding, building (1 male); running on cobblestone beach, seashore (1 male); running 

on fl attened dead grass, saltmarsh (2 females); running through grasses, old fi eld (2 females); running 

on rock, old fi eld (2 females); running in grass (2 females); running through reindeer lichen (1 male); 

running through grass near vegetable garden (4 males, 3 females); running in vegetable garden (1 

female); running on soil, vegetable garden (1 male); running on lawn (1 female); running on lawn 

with egg sac (1 female); running on lawn (1 male); running on brick walk (1 female); under rock, 

cobble beach, seashore (1 female); n. d., old fi eld (2 females).

Regional Distribution: ME (Emerton 1909, as Pardosa diffusa Emerton); NH, VT, MA, CT, QC, 

NB, NS, NF, LB (Dondale and Redner 1987, 1990).

Pardosa saxatilis (Hentz, 1844)

Taxonomy: Dondale and Redner (1984, 1990); Kaston (1981); Paquin and Dupérré (2003); Vogel 

(2004).

Record: M01 (1 male). N = 1 adult male.

Method: search (1 male).

Month: June (1 male).

Habitat: on wall of house (1 male).

Regional Distribution: ME (Bryant 1908, as Pardosa minima (Keyserling) Montgomery); NH, VT, 

MA, CT, RI, NY, ON, QC, NS, PE (Dondale and Redner 1984, 1990).

Pardosa xerampelina (Keyserling, 1877)

Taxonomy: Dondale and Redner (1986, 1990); Kaston (1981); Paquin and Dupérré (2003); Vogel 

(2004).

Records: M01 (40 males, 74 females); M06 (1 female); M07 (3 males, 4 females); M11 (3 females); 

M12 (2 females); M14 (2 males, 10 females); M16 (1 male); M22 (18 females); M24 (1 male, 1 

female); M25 (2 females); M26 (5 females); M47 (1 female); M51 (2 females). N = 170 adults; 47 

males, 123 females.

Methods: pitfall traps (17 males, 41 females); search (30 males, 81 females); sweep net (1 female).

Months: April (5 males, 1 female); May (24 males, 13 females); May-June (3 males, 2 females); June 

(15 males, 61 females); June-July (6 females); July (28 females); July-August (3 females); August 

(5 females); September (4 females).

Habitats: litter, Amelanchier-Rubus stand (7 males, 1 female); litter, seashore-backshore (6 males, 

3 females); litter, deciduous-coniferous woodland (2 females); litter, saltmarsh (1 female); litter, 

Kalmia-Vaccinium heath (3 males, 4 females); ground, gravel pit, conifer-mixed hardwood (2 females); 

litter, freshwater marsh edge (2 females); litter, bigtooth aspen stand (18 females); litter, red maple 
101

sapling stand (1 male, 1 female); litter, white (paper) birch stand (2 females); litter, saltmarsh edge 

(5 females); sweeping grasses along freshwater stream, riparian (1 female); in or on compost pile 

(crib) (9 females); running in compost pile (2 females); in woodpile (1 male, 5 females); in fl ower 

garden with egg sac (1 female); in vegetable garden (2 males, 2 females); on ground in vegetable 

garden (4 females); on ground and boards in vegetable garden (2 males, 3 females); on lettuce leaf in 

vegetable garden (1 female); running on soil or straw in vegetable garden (7 males, 6 females); in and 

around beds in vegetable garden (2 males); running on pile of agricultural peat (1 female); running 

through grass near vegetable garden (1 female); on ground, old fi eld (1 male); running through grass, 

old fi eld (2 females); on lawn (1 female); running on lawn (1 male); running on or across gravel lane, 

mixed hardwood-conifer (1 male, 2 females); among hot rocks, seashore (1 female); on cobble beach, 

seashore (1 female); on pebbly beach in grasses and dead rockweed, seashore (1 male, 1 female); 

running on beach, seashore (1 female); under rock with egg sac, seashore (1 female); on seashore 

ledges (1 male); on ground in sand-rock pit, mixed hardwood-conifer (6 females); on ground along 

forest trail, mixed hardwood-conifer (2 males, 3 females); on ground, vernal pool, mixed hardwoodconifer (1 female); on woods road, mixed conifer opening (2 females); on ground in gravel pit, mixed 

conifer-hardwood (1 female); on ground, freshwater pond edge (1 female); on grass, freshwater pond 

edge (1 female); on ground, old fi eld edge (2 males); in garage, house (1 female); in cellar entryway, 

house (1 female); on cellar door with egg sac, house (1 female); on cellar window, house (1 male); 

on outside walls (shingled) of house (2 males, 6 females); on porch, house (1 female); on outside of 

screened window, house (1 female); at base of house foundation (2 females); around house foundation 

(1 female); on house foundation (1 female); on patio, house (1 male); running on stone patio (1 

female); under umbrella base on patio, house (1 female); on deck, house (1 male); running across 

deck (1 male); fell, dropped, or drowned in outdoor hot tub (Jacuzzi) (3 females); drowned in water 

bucket (1 female); under old board, prey of Xysticus emertoni (1 male).

Regional Distribution: ME (Bryant 1908, as Pardosa tachypoda (Thorell) Emerton); ON, QC, NB, 

NS, NF, LB (Dondale and Redner 1990); CT (Kaston 1981); NY (Crosby and Bishop 1928).

Pirata cantralli Wallace & Exline, 1978

Taxonomy: Wallace and Exline (1978); Dondale and Redner (1990); Paquin and Dupérré (2003).

Records: M06 (3 females); M12 (3 females); M55 (1 female); M56 (4 females). N = 11 adult females.

Methods: litter condenser-Berlese funnels (3 females); litter condenser-hand sorted (2 females); pitfall 

traps (6 females).

Months: June (1 female); August (1 female); August-September (1 female); September (8 females).

Habitats: litter, saltmarsh (3 females); litter, freshwater marsh edge (3 females); litter, freshwater 

streamside (1 female); litter, salt meadow (4 females).

Regional Distribution: ME (Jennings et al. 1988); ON (Wallace and Exline 1978); QC (Bélanger 

and Hutchinson 1992); NB (Dondale and Redner 1990).

Note: Females of Pirata cantralli can be distinguished from those of P. insularis Emerton, 1885 by 

differences in carapace width-length ratios and coloration pattern (Jennings, unpubl.).
102

Pirata insularis Emerton, 1885

Taxonomy: Wallace and Exline (1978); Kaston (1981); Dondale and Redner (1990); Paquin and 

Dupérré (2003).

Records: M06 (3 males, 1 female); M12 (14 males, 7 females). N = 25 adults; 17 males, 8 females.

Method: pitfall traps (17 males, 8 females).

Months: June (14 males, 3 females); June-July (2 males); July-August (1 male, 1 female); August 

(3 females); August-September (1 female).

Habitats: litter, saltmarsh (3 males, 1 female); freshwater marsh edge (14 males, 7 females).

Regional Distribution: ME, NH, VT, MA, CT, NY, ON, QC (Wallace and Exline 1978); NB, NS, 

NF (Dondale and Redner 1990).

Note: Emerton (1920) included a record of this species from Kettle Rapids, Maine which, most 

likely, is an error. Elsewhere in the same publication he refers to Kettle Rapids, Manitoba.

Pirata minutus Emerton, 1885

Taxonomy: Wallace and Exline (1978); Kaston (1981); Dondale and Redner (1990); Paquin and 

Dupérré (2003).

Records: M01 (1 male, 1 female); M05 (1 female); M06 (2 males, 8 females); M07 (2 males); M24 

(1 male). N = 16 adults; 6 males, 10 females.

Methods: pitfall traps (6 males, 9 females); sweep net (1 female).

Months: June (2 males); July (4 males, 3 females); August (6 females); September (1 female).

Habitats: litter, edge of old fi eld and alder swamp (1 female); litter, seashore-backshore (1 male); 

litter, saltmarsh (2 males, 8 females); litter, Kalmia-Vaccinium heath (2 males); litter, red maple 

sapling stand (1 male); sweeping vegetation, old fi eld (1 female).

Regional Distribution: ME, NH, VT, MA, CT, NY, ON, NB, NS (Wallace and Exline 1978); QC 

(Bélanger and Hutchinson 1992); NF (Dondale and Redner 1990).

Pirata piraticus (Clerck, 1757)

Taxonomy: Wallace and Exline (1978); Kaston (1981, as Pirata piratica (Olivier)); Dondale and 

Redner (1990); Paquin and Dupérré (2003).

Records: M01 (2 males, 1 female); M06 (10 males, 9 females); M12 (1 male, 1 female); M26 

(2 males); M33 (4 males, 1 female). N = 31 adults; 19 males, 12 females.

Methods: aquatic pitfall traps (7 males, 3 females); pitfall traps (12 males, 9 females).
103

Months: May (4 males, 6 females); June (7 males, 1 female); July (8 males, 4 females); August 

(1 female).

Habitats: water surface, freshwater pond near shore (2 males, 1 female); water surface, beaver pond 

(1 male, 1 female); water surface, freshwater marsh (4 males, 1 female); litter, saltmarsh (10 males, 

9 females); litter, saltmarsh edge (2 males).

Regional Distribution: ME (Procter 1938, as Pardosa prodigiosa Keyserling (febriculosa Becker)); 

MA, CT, NY, ON, NB, QC (Wallace and Exline 1978); NS, NF (Dondale and Redner 1990).

*Pirata sp. (nr. montanus Emerton, 1885)

Taxonomy: New species.

Records: M01 (11 males); M07 (2 males); M24 (1 male); M25 (1 male); M27 (1 male). N = 16 

adult males.

Method: pitfall traps (16 males).

Months: June (5 males); June-July (8 males); July (3 males).

Habitats: litter, mixed deciduous-coniferous woodland (7 males); litter, seashore-backshore (4 males); 

litter, Kalmia-Vaccinium heath (2 males); litter, red maple sapling stand (1 male); litter, white (paper) 

birch stand (1 male); litter, red maple-white birch stand (1 male).

Regional Distribution: ME (this study); NY (Cokendolpher, unpubl.).

Note: This species resembles Pirata montanus Emerton, 1885 but differs morphologically in 

structures of the adult genitalia and phenotypically in coloration pattern. Specimens collected in 

Maine were forwarded to James C. Cokendolpher for further study and analysis.

Schizocosa communis (Emerton, 1885)

Taxonomy: Dondale and Redner (1978a, 1990); Kaston (1981); Paquin and Dupérré (2003).

Records: M40 (2 females). N = 2 adult females.

Method: search (2 females).

Month: July (2 females).

Habitat: under rocks, lowbush blueberry fi eld (2 females).

Regional Distribution: ME, NH, VT, MA, CT, NY, ON, QC, NS (Dondale and Redner 1978a, 

1990).
104

Schizocosa crassipalpata Roewer, 1951

Taxonomy: Dondale and Redner (1978a, 1990); Kaston (1981); Paquin and Dupérré (2003).

Record: M26 (1 male). N = 1 adult male.

Method: pitfall trap (1 male).

Month: June (1 male).

Habitat: litter, saltmarsh edge (1 male).

Regional Distribution: ME (Emerton 1909, as Lycosa crassipalpis Emerton); NH, VT, MA, NY, ON, 

QC, NB, NS (Dondale and Redner 1978a, 1990).

Schizocosa saltatrix (Hentz, 1844)

Taxonomy: Dondale and Redner (1978a, 1990); Kaston (1981); Paquin and Dupérré (2003).

Records: M06 (1 female); M12 (1 female); M22 (6 females); M25 (1 male, 2 females); M26 (1 

female); M27 (1 male). N = 13 adults; 2 males, 11 females.

Method: pitfall traps (2 males, 11 females).

Months: May (1 male); May-June (1 male); June (2 females); June-July (3 females); July (5 females); 

July-August (1 female).

Habitats: litter, saltmarsh (1 female); litter, freshwater marsh edge (1 female); litter, bigtooth aspen 

stand (6 females); litter, white (paper) birch stand (1 male, 2 females); litter, saltmarsh edge (1 female); 

litter, red maple-white (paper) birch stand (1 male).

Regional Distribution: ME, NH, VT, MA, CT, NY, ON, QC, NS (Dondale and Redner 1978a, 

1990).

Trabeops aurantiacus (Emerton, 1885)

Taxonomy: Russell-Smith (1982); Kaston (1981, as Trabea aurantiaca (Emerton)); Dondale and 

Redner (1990).

Records: M12 (2 males); M22 (1 male); M25 (6 males); M26 (2 males); M27 (1 female). N = 12 

adults; 11 males, 1 female.

Method: pitfall traps (11 males, 1 female).

Months: June (7 males, 1 female); June-July (1 male); July (3 males).
105

Habitats: litter, freshwater marsh edge (2 males); litter, bigtooth aspen stand (1 male); litter, white 

(paper) birch stand (6 males); litter, saltmarsh edge (2 males); litter, red maple-white (paper) birch 

stand (1 female).

Regional Distribution: ME, MA, CT, NY, NS (Dondale and Redner 1990).

Trochosa ruricola (De Geer, 1778)

Taxonomy: Roberts (1985); Paquin and Dupérré (2003).

Records: M01 (36 males, 16 females); M04 (1 female); M06 (2 males); M11 (1 female); M15 (1 

female); M26 (7 males, 2 females). N = 66 adults; 45 males, 21 females.

Methods: pitfall traps (33 males, 8 females); search (12 males, 12 females); sweep net (1 female).

Months: February (1 male); April (2 males); May (2 males, 3 females); May-June (1 male); June 

(18 males, 4 females); June-July (3 males); July (14 males, 8 females); July-August (1 male); August 

(1 male, 3 females); September (1 male, 2 females); October (1 male, 1 female).

Habitats: litter, seashore-backshore (24 males, 4 females); litter, coastal red spruce stand (1 female); 

litter, saltmarsh (2 males); ground, gravel pit in conifer-mixed hardwood (1 female); litter, saltmarsh 

edge (7 males, 2 females); sweeping grasses, saltmarsh (1 female); running through, on, or at edge 

of lawn (1 male, 3 females); running in grass (1 male, 1 female); in grass on lawn (1 male); in or on 

compost pile (bin) (1 male, 1 female); running across garden (1 female); under wood chips in garden 

(1 female); in fl ower garden (1 male); under boards in old fi eld, with prey (1 female); on rock in old 

fi eld (1 female); on wall or side of house (2 males); around foundation, house (1 female); in kitchen 

sink, house (1 male, 1 female); in bathtub, house (2 males); on bedroom window ledge, house 

(1 male); on porch (dead), house (1 female); in bucket of water, dead (1 male).

Regional Distribution: ME (Jennings, unpubl.); MA (Edwards 1993); QC (Paquin and Dupérré 

2003).

Note: This invasive species was fi rst reported in New England by Edwards (1993). It has since been 

found as far west as Illinois (Prentice 2001).

Trochosa terricola Thorell, 1856

Taxonomy: Brady (1979); Kaston (1981); Dondale and Redner (1990); Paquin and Dupérré (2003).

Records: M01 (20 males, 19 females); M04 (3 males, 5 females); M06 (11 males, 9 females); M07 

(4 males, 4 females); M12 (1 female); M15 (1 female); M24 (8 males, 4 females); M25 (4 males, 6 

females); M26 (25 males, 25 females); M27 (2 males, 3 females); M35 (1 female); M54 (1 male). N = 

156 adults; 78 males, 78 females.

Methods: litter condenser-Berlese funnel (1 male); pitfall traps (72 males, 63 females); search 

(4 males, 15 females); sweep net (1 male).
106

Months: April (2 males); May (37 males, 16 females); May-June (14 males, 6 females); June (12 

males, 5 females); June-July (2 males, 6 females); July (2 males, 23 females); July-August (10 females); 

August (5 males, 5 females); August-September (1 male, 3 females); September (3 males, 4 females).

Habitats: litter, Betula-Acer stand (2 males, 5 females); litter, Amelanchier-Rubus stand (3 males); 

litter, edge of old fi eld and alder swamp (4 males, 2 females); litter, seashore-backshore (7 males); 

litter, coastal red spruce stand (3 males, 5 females); litter, saltmarsh (10 males, 9 females); litter, 

Kalmia-Vaccinium heath (4 males, 4 females); litter, freshwater marsh edge (1 female); litter, red 

maple sapling stand (8 males, 4 females); litter, white (paper) birch stand (4 males, 6 females); litter, 

saltmarsh edge (25 males, 24 females); litter, red maple-white birch stand (2 males, 3 females); 

sifted litter, coastal red spruce stand (1 male); sweeping grasses, saltmarsh (1 male); in high saltgrass 

at shore, seashore-backshore (1 female); in vegetable garden (1 male, 1 female); in compost pile (2 

females); in low plants, lawn edge (1 female); on ground, old fi eld (1 female); on ground, saltmarsh 

edge (1 female); on ground in small burrow among marsh debris, brackish marsh (1 female); on 

house foundation (1 male); running on cobble beach, seashore (1 female); running on gravel lane 

(1 female); running through leaves, gravel lane edge (1 male); running on stone path (1 male); 

running in fl ower garden (1 female); running across bedroom fl oor, house (1 female); under bark of 

Betula papyrifera (1 female); under litter, upper wrack line, saltmarsh (1 female); under chipped bark 

in garden (1 female).

Regional Distribution: ME (Bryant 1908, as Lycosa pratensis Emerton); NH, VT, MA, RI, CT, NY, 

ON, QC, NB, NS, PE, NF, LB (Brady 1979; Dondale and Redner 1990).
107

FAMILY PISAURIDAE

The pisaurids or nursery-web spiders are among the largest spiders in Maine. Some pisaurids are 

semiaquatic, living on or near bodies of water. They can pursue and capture prey, including small 

minnows and tadpoles, among submerged vegetation of ponds, lakes, and rivers (Gertsch 1979), but 

also forage among rocks and vegetation along shorelines. Many kinds of aquatic insects are included 

among the prey captured and eaten by these large spiders. Unlike female lycosids that transport their 

egg sacs attached to the spinnerets, female pisaurids carry their egg sacs slung beneath their bodies. 

The sacs are held in place by the female’s chelicerae and pedipalps. Before the young spiderlings 

emerge, the female spins a tent-like web in which the egg sac is enclosed. The female remains nearby 

until the young spiderlings emerge and disperse; hence, the common name “nursery-web spiders.”

Although closely related to the Lycosidae, there are distinct morphological differences. The eyes are 

arranged in two rows, not three; the posterior row of eyes is strongly recurved and well separated from 

the anterior row. None of the eyes are conspicuously enlarged; the carapace is somewhat fl attened. 

Carico (2005) provides additional information on family characteristics, natural and taxonomic 

histories, and a key to the genera of North America north of Mexico.

Carico (1972, 1973) revised the Nearctic species of Pisaurina and Dolomedes, the two genera most 

likely to be encountered in Maine. Dondale and Redner (1990) also provide identifi cation keys, 

descriptions, and illustrations for pisaurids of the northern region. The Pisauridae in Maine are 

represented by two genera and fi ve species; in Milbridge, this family is represented by only one genus 

and species. Most likely additional species of Dolomedes and at least one species of Pisaurina inhabit 

this coastal region, but remain to be detected and specimens collected.

Dolomedes striatus Giebel, 1869

Taxonomy: Carico (1973); Dondale and Redner (1990); Paquin and Dupérré (2003).

Record: M01 (1 male). N = 1 adult male.

Method: pitfall trap (1 male).

Month: June (1 male).

Habitat: litter, seashore-backshore (1 male).

Regional Distribution: ME (Dondale and Redner 1990); MA, CT, NY, ON, PQ, NS, LB (Carico 

1973); NB (Dondale and Redner 1990).
108

FAMILY LIOCRANIDAE

The liocranids were formerly assigned to the subfamily Liocraninae of the family Clubionidae. 

Lehtinen (1967) elevated this subfamily to full family status and transferred some of the genera to 

other families. Like the clubionids, liocranids are hunters that do not rely on webs for prey capture. 

Species of Agroeca inhabit moist litter of fi elds, meadows, bogs, and forests (Dondale and Redner 

1982). Unlike the Clubionidae, the liocranids do not spin a retreat.

Structurally, members of this family differ from clubionids by having the palp-coxal lobes straight 

or convex along their lateral margins. The metatarsi of legs I and II have long, paired ventral spines. 

Three dorsal macroseta are present on femur I of Agroeca; species of this genus also have a dense 

cluster of long setae on the anterior margin of the abdomen. Ubick and Richman (2005b) provide 

additional family-character descriptors, notes on natural and taxonomic histories, and a key to the 

liocranid genera of North America.

With the recent removal of Phrurotimpus and Scotinella to the family Corinnidae by Bosselaers and 

Jocqué 2002, the liocranid fauna in Maine is now represented by only one genus (Agroeca) and two 

species. This genus and both species have been found in Milbridge. 

Agroeca ornata Banks, 1892

Taxonomy: Kaston (1938a, 1981); Dondale and Redner (1982); Paquin and Dupérré (2003).

Records: M01 (2 males, 2 females); M04 (7 males); M22 (2 females); M24 (8 males, 1 female); M25 

(6 males); M27 (8 males, 2 females). N = 38 adults; 31 males, 7 females.

Method: pitfall traps (31 males, 7 females).

Months: May (12 males, 1 female); May-June (8 males); June (10 males, 3 females); June-July 

(1 male, 1 female); July-August (1 female); August (1 female).

Habitats: forest-fl oor litter, Betula-Acer stand (1 female); litter, deciduous-coniferous woodland 

(2 males, 1 female); forest-fl oor litter, coastal red spruce stand (7 males); forest-fl oor litter, bigtooth 

aspen stand (2 females); forest-fl oor litter, red maple sapling stand (8 males, 1 female); forest-fl oor 

litter, white (paper) birch stand (6 males); forest-fl oor litter, mixed red maple-white birch stand (8 

males, 2 females).

Regional Distribution: ME, MA, CT, NH, VT, NY, ON, QC, NB, NS (Dondale and Redner 

1982).

Agroeca pratensis Emerton, 1890

Taxonomy: Kaston (1938a, 1981); Dondale and Redner (1982); Paquin and Dupérré (2003).

Records: M01 (1 male); M22 (2 males); M25 (1 male); M26 (5 females). N = 9 adults, 4 males, 

5 females.

Methods: pitfall traps (3 males, 5 females); search (1 male).
109

Months: May (1 male, 1 female); June-July (1 female); July (3 females); August-September (1 male); 

September (1 male); November (1 male).

Habitats: forest-fl oor litter, bigtooth aspen stand (2 males); forest-fl oor litter, white (paper) birch 

stand (1 male); litter, saltmarsh edge (5 females); on patio edge near fl ower garden (1 male).

Regional Distribution: ME (Kaston 1938a); NH, VT, MA, CT, RI, NY, ON, QC, NB, NS 

(Dondale and Redner 1982).

FAMILY CLUBIONIDAE

The clubionids or sac spiders are small tubular-shaped spiders whose bodies are sparsely covered 

with setae. They are usually creamy yellow, but may have brown markings on the abdomen. They 

are nocturnal hunters and spend the day in tubular retreats or sacs; hence, the common name. In 

addition to temporary retreats, these sacs also serve as pre-nuptial chambers and nests for egg-laying. 

The female usually remains in the nest until the young spiderlings emerge.

Most clubionids are swift runners that move rapidly when disturbed. They seize their prey with stout, 

toothed chelicerae. Some species inhabit grasses and shrubs; others are found in bogs or on foliage of 

trees. Some of the tree-inhabiting species can be collected by jarring limbs and foliage over a beating 

cloth.

Members of the Clubionidae are two-clawed spiders with a dense brush of setae or claw tuft on the 

pretarsus. Ventrally, the distal leg segments bear a brush of setae called scopulae. From one to three 

macrosetae are present on the prolateral surface of femur I. The eyes are arranged in two transverse 

rows near the anterior margin of the carapace. The eye rows may be straight, recurved, or procurved. 

The anterior spinnerets are close together and not heavily sclerotized; the palp-coxal lobes lack a 

depression on the ventral surface. Additional character descriptors, notes on natural history, and a key 

to North American genera of Clubionidae are provided by Richman and Ubick (2005). Descriptions 

and illustrations of Nearctic species of Clubionidae are provided by Edwards (1958), Dondale and 

Redner (1982), and Paquin and Dupérré (2003).

In Maine, the Clubionidae are represented by 2 genera and 20 species. Thirteen species of the genus 

Clubiona are found in Milbridge.

Clubiona bishopi Edwards, 1958

Taxonomy: Edwards (1958); Dondale and Redner (1982); Paquin and Dupérré (2003).

Records: M01 (1 male, 1 female); M06 (1 male); M23 (3 males, 1 female). N = 7 adults; 5 males, 

2 females.

Methods: pitfall trap (1 male); sweep net (4 males, 2 females).

Months: June (1 male, 1 female); July (3 males, 1 female); September (1 male).
110

Habitats: sweeping vegetation, freshwater pond edge (1 male, 1 female); sweeping Vaccinium sp., 

mixed conifer-hardwood, island (1 male, 1 female); sweeping grassy area near cabin, mixed conifer 

opening, island (2 males); litter, saltmarsh (1 male).

Regional Distribution: ME (Edwards 1958); NY, ON, QC, NB, NS (Dondale and Redner 1982).

Clubiona bryantae Gertsch, 1941

Taxonomy: Edwards (1958); Dondale and Redner (1982); Paquin and Dupérré (2003).

Records: M01 (1 female); M12 (2 females); M33 (1 female). N = 4 adult females.

Methods: pitfall trap (1 female); search (1 female); sweep net (2 females).

Months: June (1 female); July (2 females); August (1 female).

Habitats: sweeping old fi eld vegetation (1 female); sweeping grasses, freshwater marsh edge 

(1 female); litter, freshwater marsh edge (1 female); in folded leaf of leatherleaf, freshwater marsh 

(1 female).

Regional Distribution: ME, MA, NY (Edwards 1958); ON, QC, NB, NS, NF (Dondale and 

Redner 1982). 

Clubiona canadensis Emerton, 1890

Taxonomy: Edwards (1958); Dondale and Redner (1982); Paquin and Dupérré (2003).

Records: M01 (3 males, 10 females); M04 (2 males, 3 females); M06 (1 female); M15 (1 female); 

M23 (2 females); M29 (1 female); M30 (1 female); M42 (1 female); M55 (1 female). N = 26 adults; 

5 males, 21 females.

Methods: beating cloth (1 female); brush-tree bole net (1 female); pitfall traps (5 males, 4 females); 

search (10 females); sweep net (3 females); n. d. (2 females).

Months: June (1 male, 5 females); July (3 males, 9 females); July-August (1 male); August 

(5 females); September (2 females).

Habitats: litter, Betula-Acer stand (3 males); litter, coastal red spruce stand (2 males, 1 female); litter, 

deciduous-coniferous woodland (2 females); litter, Amelanchier-Rubus stand (1 female); sweeping 

shrubby vegetation, coastal spruce-mixed hardwood (1 female); sweeping understory, spruce-mixed 

hardwood (1 female); sweeping grasses, forbs, ferns, and rushes, roadside alder-aspen edge (1 female); 

brushing white (paper) birch bark (1 female); beating red spruce foliage, near freshwater stream 

(1 female); in silk retreat, folded wild-raisin leaf (1 female); in silk retreat, folded goldenrod leaf, 

freshwater pond (1 female); in silk retreat between Amelanchier leaves, Amelanchier-Rubus stand 

(1 female); in silk retreat under dead bark of apple (1 female); in silk retreat, meadowsweet leaf, old 

fi eld (1 female); in silk retreat with egg sac, folded aster leaf, old fi eld (1 female); on balsam fi r snag 

(1 female); under rocks in silk retreat, open ledges-mixed conifer, island (1 female); n. d., cedar, 
111

mixed conifer-hardwood, island (1 female); n. d., cherry sp., mixed conifer-hardwood, island (1 

female); in silk retreat, mailbox (1 female); in mailbox (1 female).

Regional Distribution: ME (Emerton 1914); NH, VT, CT, NY, ON, QC, NB, NS, NF, LB 

(Edwards 1958).

Clubiona johnsoni Gertsch, 1941

Taxonomy: Edwards (1958); Dondale and Redner (1982); Paquin and Dupérré (2003).

Records: M05 (1 male); M12 (1 female); M26 (2 males, 3 females); M35 (1 female); M56 (2 males, 

7 females). N = 17 adults; 5 males, 12 females.

Methods: litter condenser-Berlese funnel (1 male, 3 females); litter condenser-hand sorted (1 male, 

4 females); pitfall traps (2 males, 3 females); search (1 female); sweep net (1 female); n. d. (1 male).

Months: June (1 female); July (1 male, 2 females); July-August (1 male, 2 females); August (1 male); 

September (2 males, 7 females). 

Habitats: sifted litter (grasses, forbs, tidal debris), salt meadow (2 males, 7 females); litter, saltmarsh 

edge (2 males, 3 females); sweeping shrubs and grasses, freshwater marsh (1 female); in silk retreat, 

dry forb stem, brackish marsh (1 female); old fi eld (1 male).

Regional Distribution: ME (Collins et al. 1996); NH, VT, MA, CT, RI, NY, ON, NS (Edwards 

1958); QC, NB (Dondale and Redner 1982).

Clubiona kastoni Gertsch, 1941

Taxonomy: Edwards (1958); Dondale and Redner (1982); Paquin and Dupérré (2003).

Records: M06 (1 male); M12 (3 males, 6 females); M14 (2 females). N = 12 adults; 4 males, 

8 females.

Methods: pitfall traps (4 males, 6 females); sweep net (2 females).

Months: July (2 females); August (4 males, 5 females); September (1 female).

Habitats: litter, freshwater marsh edge (3 males, 6 females); litter, saltmarsh (1 male); sweeping 

grasses and forbs, mixed hardwood-conifer (2 females).

Regional Distribution: ME (Gertsch 1941); NH, VT, MA, CT, NY, ON, NB (Edwards 1958); QC, 

NS (Dondale and Redner 1982).

*Clubiona littoralis Banks, 1895

Taxonomy: Edwards (1958); Dondale and Redner (1982).

Records: M05 (2 males, 2 females); M15 (1 female). N = 5 adults; 2 males, 3 females.
112

Methods: sifted litter-hand sorted (2 males, 2 females); sweep net (1 female)

Months: July (1 female); August (2 males, 2 females).

Habitats: sifted dried seaweed, old fi eld-brackish marsh edge (2 males, 2 females); sweeping grasses 

and sedges, saltmarsh (1 female).

Regional Distribution: ME (this study); MA, NY, ON (Edwards 1958); NS (Dondale and Redner 

1982).

*Clubiona maritima L. Koch, 1867

Taxonomy: Edwards (1958); Dondale and Redner (1982); Paquin and Dupérré (2003).

Record: M12 (1 female). N = 1 adult female.

Method: search (1 female).

Month: August (1 female).

Habitat: in silk retreat (with spiderlings), folded grass stem, freshwater marsh (1 female).

Regional Distribution: ME (this study); MA, CT, NY, ON (Edwards 1958); QC (Bélanger and 

Hutchinson 1992).

Clubiona moesta Banks, 1896

Taxonomy: Edwards (1958); Dondale and Redner (1982); Paquin and Dupérré (2003).

Record: M01 (1 female). N = 1 adult female.

Method: search (1 female).

Month: July (1 female).

Habitat: in alders along seashore, seashore-backshore (1 female).

Regional Distribution: ME (Procter 1938, as Clubiona emertoni Petrunkevitch (pusilla Emerton)); 

NH, MA, NY, ON (Edwards 1958); VT, QC, NB, NS (Dondale and Redner 1982).

Clubiona norvegica Strand, 1900

Taxonomy: Edwards (1958); Dondale and Redner (1982); Paquin and Dupérré (2003).

Records: M12 (3 females); M15 (2 females); M33 (5 males, 5 females). N = 15 adults; 5 males, 10 

females.

Method: sweep net (5 males, 10 females).
113

Months: June (2 males, 5 females); July (3 males, 5 females).

Habitats: sweeping grasses and shrubs (Spiraea-Kalmia), freshwater marsh (3 females); sweeping 

grasses and sedges, freshwater marsh (5 males, 5 females); sweeping grasses, saltmarsh (2 females).

Regional Distribution: ME (Longcore et al., unpubl.); NY, ON, NF, LB (Edwards 1958); QC 

(Dondale and Redner 1982).

Clubiona obesa Hentz, 1847

Taxonomy: Edwards (1958); Dondale and Redner (1982); Paquin and Dupérré (2003).

Record: M01 (1 female). N = 1 adult female.

Method: search (1 female).

Month: June (1 female).

Habitat: in silk retreat on willow leaf, freshwater pond edge (1 female).

Regional Distribution: ME, NH, VT, MA, CT, RI, NY, ON (Edwards 1958); QC, NS, NF 

(Dondale and Redner 1982).

Clubiona pygmaea Banks, 1892

Taxonomy: Edwards (1958); Dondale and Redner (1982); Paquin and Dupérré (2003).

Record: M35 (1 male). N = 1 adult male.

Method: sweep net (1 male).

Month: June (1 male).

Habitat: sweeping grasses and sedges, brackish marsh (1 male).

Regional Distribution: ME (Bryant 1908, as Clubiona minuta Emerton); MA, CT, NY, ON, QC 

(Edwards 1958).

Clubiona riparia L. Koch, 1866

Taxonomy: Edwards (1958); Dondale and Redner (1982); Paquin and Dupérré (2003).

Records: M01 (4 females); M05 (3 males, 3 females); M06 (3 males, 26 females); M08 (2 males); 

M12 (11 males, 18 females); M17 (8 females); M23 (1 male, 4 females); M33 (9 males, 15 females); 

M34 (1 female); M35 (2 males, 8 females); M47 (1 female); M58 (1 male, 3 females). N = 123 

adults; 32 males, 91 females.
114

Methods: pitfall trap (1 female); search (26 females); sweep net (32 males, 63 females); n. d. 

(1 female).

Months: May (5 males, 2 females); June (8 males, 28 females); July (19 males, 30 females); August 

(30 females); September (1 female).

Habitats: sweeping vegetation (Juncus-Spartina-shrubs), saltmarshes (3 males, 27 females); sweeping 

grasses, sedges, shrubs (Spiraea-Kalmia), freshwater marshes (20 males, 16 females); sweeping grasses, 

forbs, rushes (Juncus), seashore brackish marsh (3 males, 10 females); sweeping grasses and forbs, 

old fi eld (3 males, 4 females); sweeping blue-joint grass, seashore-backshore (1 male, 4 females); 

sweeping upper beach grasses, forbs, seaside-pea, and shrubs, seashore-backshore (2 males); sweeping 

vegetation, dry stream bed (1 female); sweeping marsh vegetation near freshwater stream (1 female); 

in silk retreats, folded marsh grasses (Spartina sp.), freshwater marsh (13 females); in silk retreat, 

folded marsh grass, brackish marsh (1 female); in silk retreats, folded Spiraea leaves, freshwater marsh 

(6 females); in silk retreats, folded saltmarsh grasses, saltmarshes (3 females); in silk retreats, folded 

stems of timothy grass (2 females); in silk retreat, folded Spiraea leaf, old fi eld (1 female); litter, 

saltmarsh (1 female); n. d., saltmarsh (1 female).

Regional Distribution: ME (Bryant 1908); NH, MA, CT, NY, ON, QC, NB, NS (Edwards 1958); 

NF (Dondale and Redner 1982).

Clubiona trivialis C. L. Koch, 1843

Taxonomy: Edwards (1958); Dondale and Redner (1982); Paquin and Dupérré (2003).

Records: M01 (4 males, 10 females); M02 (1 female); M04 (1 male, 1 female); M06 (2 males); M10 

(1 male, 3 females); M15 (1 male, 2 females); M29 (1 male); M48 (6 males, 1 female); M49 (1 male). 

N = 35 adults; 17 males, 18 females.

Method: beating cloth (17 males, 18 females).

Months: May (3 males, 11 females); June (1 male, 4 females); July (3 males); August (4 males, 

2 females); September (6 males, 1 female).

Habitats: beating red spruce foliage, seashore-backshore (8 males, 3 females); beating red spruce 

foliage (saplings), conifer-mixed hardwood (1 male, 3 females); beating red spruce foliage, saltmarsh 

edge (1 male, 2 females); beating red spruce foliage, old fi eld edge (1 female); beating red spruce 

foliage, roadside (1 male); beating white spruce foliage, saltmarsh edge (2 males); beating young 

balsam fi r foliage, mixed conifer (1 female); beating spruces, old fi eld edge (4 males, 7 females); 

beating spruce, seashore (1 female).

Regional Distribution: ME (Procter 1946, as Clubiona obtusa Emerton); NY, ON, LB, NF 

(Edwards 1958); QC, NB, NS (Dondale and Redner 1982).
115

FAMILY CORINNIDAE

Formerly a subfamily of the Clubionidae, this family has now been elevated to full family status. It 

contains genera previously assigned to one or more subfamilies of Clubionidae and subsequently to 

other spider families such as the Gnaphosidae and Liocranidae. The genera and species in our region 

are generally antlike in habitus and movements and are usually associated with ground litter of old 

fi elds, meadows, bogs, and forests. Species of Scotinella have been found in ant nests (Dondale and 

Redner 1982). Some members have iridescent scales; others are patterned with bands of white scalelike setae on their abdomens. Most members are fast runners and swiftly regain cover when disturbed, 

or become motionless (Phrurotimpus) with their legs fl exed above an iridescent carapace, thus aiding 

in concealment. Unlike the clubionids (sensu stricto), the corinnids do not spin sacs or retreats, and 

the females abandon their egg sacs after egg-laying.

In general, the members of this family have the eyes arranged in two transverse rows; the lateral 

margins of the palp-coxal lobes lack constrictions; the legs are usually slender, with or without 

scopulae and claw tufts; femur I with or without a macroseta; and anterior abdominal margins 

lacking strong, erect setae. Ubick and Richman (2005a) provide additional family character 

descriptors, notes on natural history, and a key to the genera of Corinnidae in North America.

Reiskind (1969) revised the North and Central American species of Castianeira; the genus 

Phrurotimpus is currently under revision by colleagues at the American Museum of Natural History. 

With the recent transfer of Phrurotimpus and Scotinella from the Liocranidae to the Corinnidae (see 

Bosselaers and Jocqué 2002), the Maine fauna now consists of 4 genera and 14 species; 3 genera and 

8 species are represented in the Milbridge fauna. Dondale and Redner (1982) provide identifi cation 

keys and illustrations for the genera and species found in Maine.

Castianeira cingulata (C. L. Koch, 1841)

Taxonomy: Reiskind (1969); Kaston (1981); Dondale and Redner (1982); Paquin and Dupérré 

(2003).

Records: M01 (7 females); M22 (3 males, 1 female); M24 (6 females); M25 (6 females); M27 

(6 females). N = 29 adults; 3 males, 26 females.

Methods: pitfall traps (3 males, 25 females); search (1 female).

Months: May (1 female); June (4 females); June-July (4 females); July (13 females); July-August 

(1 female); August (3 males, 3 females).

Habitats: litter, red maple sapling stand (6 females); litter, white (paper) birch stand (6 females); 

litter, red maple-white birch stand (6 females); litter, mature Betula-Acer stand (5 females); litter, 

bigtooth aspen stand (3 males, 1 female); litter, Kalmia-Vaccinium heath (1 female); under small rock 

(1 female).

Regional Distribution: ME (Bryant 1908, as Castaneira [sic] bivittata Keyserling); NH, MA, CT, 

NY, ON (Reiskind 1969); QC, NB, NS (Dondale and Redner 1982).
116

Castianeira descripta (Hentz, 1847)

Taxonomy: Reiskind (1969); Kaston (1981); Dondale and Redner (1982); Paquin and Dupérré 

(2003).

Records: M06 (3 males, 4 females); M07 (1 male); M11 (1 female). N = 9 adults; 4 males, 5 females.

Method: pitfall traps (4 males, 5 females).

Months: July (2 males, 1 female); July-August (2 males); August (4 females).

Habitats: litter, saltmarshes (3 males, 4 females); litter, Kalmia-Vaccinium heath (1 male); gravel pit, 

conifer-mixed hardwood (1 female).

Regional Distribution: ME (Bryant 1908, as Castaneira [sic] crocata Hentz); NH, VT, MA, NY 

(Reiskind 1969); CT (Kaston 1981); ON, QC, NB, NS (Dondale and Redner 1982).

Castianeira gertschi Kaston, 1945

Taxonomy: Reiskind (1969); Kaston (1945, 1981); Dondale and Redner (1982).

Records: M06 (2 males). N = 2 adult males.

Method: pitfall traps (2 males).

Months: August-September (2 males).

Habitat: litter, saltmarsh (2 males).

Regional Distribution: ME (Jaros-Su et al., unpubl.); MA, CT, NY (Reiskind 1969); VT, ON 

(Dondale and Redner 1982). 

Castianeira longipalpa (Hentz, 1847)

Taxonomy: Reiskind (1969); Kaston (1981); Dondale and Redner (1982); Paquin and Dupérré 

(2003).

Records: M06 (1 male); M07 (1 male). N = 2 adult males.

Method: pitfall traps (2 males).

Months: August (1 male); September (1 male).

Habitats: litter, Kalmia-Vaccinium heath (1 male); litter, saltmarsh (1 male).

Regional Distribution: ME (Bryant 1908, as Castaneira [sic] pinnata (Emerton)); NH, VT, MA, 

CT, NY, ON, NS (Reiskind 1969); QC, NB (Dondale and Redner 1982).
117

Phrurotimpus alarius (Hentz, 1847)

Taxonomy: Kaston (1981); Dondale and Redner (1982); Bosselaers and Jocquè (2002); Paquin and 

Dupérré (2003).

Records: M01 (6 males, 8 females); M22 (1 male, 13 females); M24 (10 males, 4 females); M25 

(10 males, 6 females); M26 (3 males, 1 female); M27 (9 males, 5 females); M53 (1 female). N = 77 

adults; 39 males, 38 females.

Methods: litter condenser-Berlese funnel (1 female); pitfall traps (39 males, 33 females); search 

(2 females); sifted litter-hand sorted (2 females).

Months: May-June (5 males); June (18 males, 10 females); June-July (5 males, 9 females); July (10 

males, 14 females); July-August (2 females); August (3 females); August-September (1 male).

Habitats: litter, white (paper) birch stand (10 males, 6 females); litter, bigtooth aspen stand (1 male, 

13 females); litter, red maple sapling stand (10 males, 4 females); litter, red maple-white birch 

stand (9 males, 5 females); litter, saltmarsh edge (3 males, 1 female); litter, deciduous-coniferous 

woodland (2 males, 5 females); litter, Amelanchier-Rubus stand (2 males); litter, Kalmia-Vaccinium 

heath (1 male); litter, Betula-Acer stand (1 male); litter-seashore (1 female); litter, aspen-maple stand 

(1 female); in low plants at edge of lawn (1 female); drowned in bucket of water (1 female).

Regional Distribution: ME (Bryant 1908, as Micariosoma alarium (Hentz) Simon); CT (Kaston 

1981); ON, QC, NS (Dondale and Redner 1982).

Phrurotimpus borealis (Emerton, 1911)

Taxonomy: Kaston (1981); Dondale and Redner (1982); Bosselaers and Jocquè (2002); Paquin and 

Dupérré (2003).

Records: M01 (4 males, 4 females); M07 (2 females); M12 (2 females); M22 (1 male, 1 female); 

M24 (4 males, 1 female); M25 (1 male); M26 (5 males, 7 females); M27 (1 female); M28 (1 female) 

M42 (1 female). N = 35 adults; 15 males, 20 females.

Methods: litter condenser-Berlese funnel (1 female); pitfall traps (11 males, 14 females); search 

(4 males, 5 females).

Months: May (1 female); June (8 males, 1 female); June-July (3 males, 3 females); July (3 males, 11 

females); July-August (1 male, 3 females); August (1 female).

Habitats: litter, saltmarsh edge (5 males, 7 females); litter, red maple sapling stand (4 males, 1 

female); litter, bigtooth aspen stand (1 male, 1 female); litter, Kalmia-Vaccinium heath (2 females); 

litter, freshwater pond edge (2 females); litter, white (paper) birch stand (1 male); litter, red maplewhite birch stand (1 female); sifted rockweed debris, seashore (1 female); under rocks on ledges, 

red spruce-mixed conifer, island (1 female); on rock, roadside disturbed area (1 female); on fl oor in 

house (1 female), on bathroom wall (1 male), on bedroom wall (1 female); in bathtub (2 males), in 

bathroom (1 male); in tall grass under canoe, old fi eld (1 female).
118

Regional Distribution: ME, MA, NY (Kaston 1945); CT (Kaston 1981); ON, QC, NB, NS 

(Dondale and Redner 1982).

Scotinella divesta (Gertsch, 1941)

Taxonomy: Dondale and Redner (1982); Bosselaers and Jocquè (2002); Paquin and Dupérré (2003).

Records: M12 (1 female); M15 (2 females); M22 (1 female); M24 (1 female); M25 (1 female); M26 

(1 female); M27 (1 male); M42 (1 female). N = 9 adults; 1 male, 8 females.

Methods: litter condenser-Berlese funnel (1 female); pitfall traps (1 male, 5 females); search 

(1 female); sweep net (1 female).

Months: May (2 females); May-June (1 male, 1 female); June (2 females); June-July (1 female); July 

(2 females).

Habitats: sifted litter, saltmarsh (1 female); litter, saltmarsh edge (1 female); grasses, saltmarsh edge 

(1 female); litter, freshwater pond edge (1 female); litter, bigtooth aspen stand (1 female); litter, red 

maple sapling stand (1 female); litter, white (paper) birch stand (1 female); litter, red maple-white 

birch stand (1 male); under rock, mixed conifer, island (1 female).

Regional Distribution: ME (Gertsch 1941); NH, MA, NY, NB, NS (Dondale and Redner 1982).

Scotinella pugnata (Emerton, 1890)

Taxonomy: Dondale and Redner (1982); Bosselaers and Jocquè (2002); Paquin and Dupérré (2003).

Records: M12 (1 male, 1 female). N = 2 adults; 1 male, 1 female.

Method: pitfall traps (1 male, 1 female).

Month: August (1 male, 1 female).

Habitat: litter, freshwater pond edge (1 male, 1 female).

Regional Distribution: ME (Bryant 1908, as Micariosoma pugnatum (Emerton) Simon); CT 

(Kaston 1981); NY, ON, QC, NB, NS (Dondale and Redner 1982).
119

FAMILY GNAPHOSIDAE

The gnaphosids or ground spiders are hunters that capture their prey by stealth. As their common 

name implies, they live in leaf litter on the ground, and under rocks, logs, and loose bark of standing 

trees and stumps. Most are nocturnal hunters that spend the daylight hours in silken retreats; others 

hunt during the day, with some specializing on ants as food. After mating, the females lay their eggs 

in masses that are enclosed in silken sacs. The egg sacs vary in size, color, and shape; depending 

on species, the female may remain on guard with her egg sac or leave it exposed. Some species 

camoufl age their egg sacs with bits of sand, soil, or plant debris. Females of Drassodes deposit their egg 

sacs in fl imsy retreats spun under rocks and loose bark of trees and stumps; the females remain with 

the eggs until the young spiderlings emerge.

Although the gnaphosids share many characters in common with clubionids and related families 

of hunters, they can be distinguished by an oblique depression on the ventral surface of each palpcoxal lobe; anterior spinnerets cylindrical, elongate, and well sclerotized, and with their bases well 

separated; many species with irregularly shaped secondary (anterior lateral, and posterior lateral) eyes. 

Ubick (2005b) provides additional information on family-character descriptors, notes on natural 

history, and a key to the genera found in North America.

The North American species of Gnaphosidae are well known and described following the excellent 

generic revisions of Norman I. Platnick and associates at The American Museum of Natural History, 

New York. See Platnick (2007) for a list of these revisionary works published chiefl y between 1975 

and 1988. Platnick and Dondale (1992) provide descriptions, illustrations, and identifi cation keys for 

gnaphosids most apt to be found in the northeastern region. The gnaphosid fauna in Maine currently 

comprises 10 genera and 33 species; 9 genera and 17 species are represented in Milbridge.

Callilepis pluto Banks, 1896

Taxonomy: Platnick (1975); Platnick and Dondale (1992); Paquin and Dupérré (2003).

Records: M04 (2 females); M06 (1 female); M25 (3 males; 1 female); M40 (1 female); M42 

(4 females). N = 12 adults; 3 males, 9 females.

Methods: pitfall traps (3 males, 2 females); search (6 females); n. d. (1 female).

Months: June (2 males, 1 female); July (1 male, 6 females); August (2 females).

Habitats: litter, white (paper) birch (Betula papyrifera Marsh.) stand (3 males, 1 female); litter, 

saltmarsh (1 female); under rock near driveway, coastal spruce-mixed hardwood (1 female); under 

rock in silk retreat, open ledges in mixed conifer, island (1 female); under rock, mixed conifer, island 

(1 female); under ledge rock with egg sac, mixed conifer-reindeer moss, island (1 female); on ledge, 

mixed conifer, island (1 female); on ground, blueberry fi eld edge (1 female); n. d. (1 female).

Regional Distribution: ME, MA, NH, VT, CT, NY, ON, QC (Platnick 1975); NS, PE (Platnick 

and Dondale 1992).
120

Drassodes neglectus (Keyserling, 1887)

Taxonomy: Platnick and Shadab (1976); Platnick and Dondale (1992); Paquin and Dupérré (2003).

Records: M04 (1 female); M10 (1 male); M11 (1 male); M39 (6 females); M40 (1 female). N = 10 

adults; 2 males, 8 females.

Method: pitfall trap (1 male); search (1 male, 8 females).

Months: May (1 male); July (1 male, 1 female); August (7 females).

Habitats: in silk retreats under ledge rocks, maple-oak (Acer-Quercus) stand (6 females); under rock 

in silk retreat with egg sac, coastal spruce-mixed hardwood (1 female); under rocks on ledge, mixed 

conifer-hardwood (1 male); on ground, edge of blueberry fi eld (1 female); on ground in gravel pit, 

conifer-mixed hardwood (1 male).

Regional Distribution: ME (Bryant 1908); MA, NH, VT, CT, NY, ON, QC, NS (Platnick and 

Shadab 1976); NB, PE (Platnick and Dondale 1992).

Note: Emerton (1907) recorded Drassus saccatus Emerton from Long Island, Cumberland County, 

ME; however, Kaston (1981) listed Emerton’s Drassus saccatus as a junior synonym of Drassodes 

neglectus Keyserling. Platnick and Dondale (1992) list Drassus saccatus as a junior synonym of 

Drassodes saccatus (Emerton, 1890); hence, Emerton’s early record from Maine probably refers to 

saccatus instead of neglectus.

Drassyllus niger (Banks, 1896)

Taxonomy: Platnick and Shadab (1982); Platnick and Dondale (1992); Paquin and Dupérré (2003).

Records: M01 (2 females): M07 (1 male, 1 female); M12 (1 male); M22 (4 males); M25 (3 males); 

M26 (4 males, 4 females); M27 (1 male). N = 21 adults; 14 males, 7 females.

Methods: pitfall traps (14 males, 6 females); search (1 female).

Months: May-June (3 males); June (6 males, 3 females); June-July (2 males, 1 female); July (3 males, 

3 females).

Habitats: litter, saltmarsh edge (4 males, 4 females); litter, bigtooth aspen stand (4 males); litter, 

white (paper) birch stand (3 males); litter, Kalmia-Vaccinium heath (1 male, 1 female); litter, 

Amelanchier-Rubus stand (1 female); litter, freshwater marsh edge (1 male); litter, red maple-white 

(paper) birch stand (1 male); on wall of house (1 female).

Regional Distribution: ME, MA, CT, VT, ON, QC, NS (Platnick and Shadab 1982); NH (Platnick 

and Dondale 1992). 

Drassyllus socius Chamberlin, 1922

Taxonomy: Platnick and Shadab (1982); Platnick and Dondale (1992); Paquin and Dupérré (2003).
121

Records: M01 (1 male); M07 (1 male); M26 (3 males, 1 female); M42 (1 female). N = 7 adults; 

5 males, 2 females.

Methods: pitfall traps (5 males, 1 female); search (1 female).

Months: May (3 males); June (1 male, 1 female); June-July (1 male); July (1 female).

Habitats: litter, saltmarsh edge (3 males, 1 female); litter, seashore-backshore (1 male); litter, KalmiaVaccinium heath (1 male); on ledge, mixed conifer, island (1 female).

Regional Distribution: ME (Collins et al. 1996); MA, NH, ON, QC, NB, NS (Platnick and 

Shadab 1982).

Gnaphosa muscorum (L. Koch, 1866)

Taxonomy: Platnick and Shadab (1975a); Platnick and Dondale (1992); Paquin and Dupérré 

(2003).

Records: M01 (1 male, 1 female); M04 (1 female); M24 (1 male). N = 4 adults; 2 males, 2 females.

Methods: pitfall trap (1 male); search (1 male, 1 female); sweep net (1 female).

Months: May (2 males); June (1 female); July (1 female).

Habitats: under stone, old fi eld (1 male); swept from shrubby vegetation, coastal spruce-mixed 

hardwood (1 female); litter, red maple sapling stand (1 male); in bed, house (1 female).

Regional Distribution: ME (Bishop 1923, as Gnaphosa gigantes (sic.) Keyserling); MA,CT, NH, VT, 

NY, ON, QC, NS, LB (Platnick and Shadad 1975); NB, PE (Platnick and Dondale 1992).

Gnaphosa parvula Banks, 1896

Taxonomy: Platnick and Shadab (1975a); Platnick and Dondale (1992); Paquin and Dupérré 

(2003).

Records: M01 (1 male, 1 female); M06 (6 males, 3 females); M07 (1 male); M12 (3 males, 7 

females); M25 (2 males, 2 females); M26 (1 male); M27 (2 females). N = 29 adults; 14 males, 15 

females.

Methods: pitfall traps (13 males, 14 females); search (1 male, 1 female).

Months: May (2 males); May-June (2 females); June (6 males, 5 females); June-July (1 male, 

1 female); July (5 males, 3 females); August (2 females); August-September (1 female); September 

(1 female).

Habitats: litter, freshwater marsh edge (3 males, 7 females); litter, saltmarsh (6 males, 3 females); 

litter, saltmarsh edge (1 male); litter, white (paper) birch (Betula papyrifera Marsh.) stand (2 males, 
122

2 females); litter, red maple-white (paper) birch stand (2 females); litter, Kalmia-Vaccinium heath 

(1 male); in dead rockweed on gravel beach, seashore (1 male); in grass under base of lawn umbrella 

(1 female).

Regional Distribution: ME, NH, MA, CT, NY, ON, QC, NB, NS (Platnick and Shadab 1975a); 

NF (Platnick and Dondale 1992).

Haplodrassus bicornis (Emerton, 1909)

Taxonomy: Platnick and Shadab (1975b); Platnick and Dondale (1992); Paquin and Dupérré 

(2003).

Records: M07 (5 males, 1 female). N = 6 adults; 5 males, 1 female.

Method: pitfall traps (5 males, 1 female).

Months: May (4 males); June (1 male); July (1 female).

Habitat: litter, Kalmia-Vaccinium heath (5 males, 1 female).

Regional Distribution: ME (Collins et al. 1996); NH, MA, CT, NY, ON, QC (Platnick and Shadab 

1975b); NB, NS (Platnick and Dondale 1992).

Haplodrassus hiemalis (Emerton, 1909)

Taxonomy: Platnick and Shadab (1975b); Platnick and Dondale (1992); Paquin and Dupérré 

(2003).

Records: M01 (2 males, 3 females); M25 (1 female); M35 (1 female). N = 7 adults; 2 males, 

5 females.

Methods: pitfall traps (1 male, 3 females); search (1 male, 1 female); sweep net (1 female).

Months: June (2 females); July (2 males, 2 females); September (1 female).

Habitats: litter, Betula-Acer stand (1 male, 1 female); litter, Amelanchier-Rubus stand (1 female); 

litter, white (paper) birch stand (1 female); swept from brackish marsh (1 female); under plastic bag, 

outdoor woodbin (1 female); in bathtub, house (1 male).

Regional Distribution: ME, NH, VT, MA, CT, NY, ON, QC, NS, NF (Platnick and Shadab 

1975b); NB, PE (Platnick and Dondale 1992).

Haplodrassus signifer (C. L. Koch, 1839)

Taxonomy: Platnick and Shadab (1975b); Platnick and Dondale (1992); Paquin and Dupérré (2003).

Records: M04 (1 female); M07 (3 males, 1 female); M40 (1 female). N = 6 adults; 3 males, 

3 females.
123

Methods: pitfall traps (3 males, 1 female); search (2 females).

Months: May (2 males, 1 female); June (1 male); July (1 female); August (1 female).

Habitats: litter, Kalmia-Vaccinium heath (3 males, 1 female); under rocks near park drive, coastal 

spruce-mixed hardwood (1 female); under rock, blueberry fi eld (1 female).

Regional Distribution: ME (Procter 1946); NH, VT, MA, CT, NY, ON, QC, NS (Platnick and 

Shadab 1975b); PE (Platnick and Dondale 1992).

Herpyllus ecclesiasticus Hentz, 1832

Taxonomy: Platnick and Shadab (1977); Platnick and Dondale (1992); Paquin and Dupérré (2003).

Records: M01 (2 males, 1 female). N = 3 adults; 2 males, 1 female.

Method: search (2 males, 1 female).

Months: August (2 males); September (1 female).

Habitats: on screen door, house (1 male); in kitchen sink, house (1 female); in grass, lawn (1 male).

Regional Distribution: ME (Bryant 1908); MA, CT, NH, VT, NY, ON, QC, NS (Platnick and 

Shadab 1977); PE (Platnick and Dondale 1992).

*Micaria elizabethae Gertsch, 1942

Taxonomy: Platnick and Shadab (1988); Platnick and Dondale (1992).

Records: M26 (2 males). N = 2 adult males.

Method: pitfall traps (2 males).

Months: May (1 male); July-August (1 male).

Habitat: litter, saltmarsh edge (2 males).

Regional Distribution: ME (this study); MA, CT, NY, RI, ON (Platnick and Shadab 1988).

Micaria gertschi Barrows & Ivie, 1942

Taxonomy: Platnick and Shadab (1988); Platnick and Dondale (1992).

Records: M06 (1 male, 1 female). N = 2 adults; 1 male, 1 female.

Method: pitfall traps (1 male, 1 female).

Months: June (1 male); July-August (1 female).
124

Habitat: litter, saltmarsh (1 male, 1 female).

Regional Distribution: ME, MA, CT, NH, VT, NY, ON, NB, NS (Platnick and Shadab 1988).

Micaria pulicaria (Sundevall, 1831)

Taxonomy: Platnick and Shadab (1988); Platnick and Dondale (1992); Paquin and Dupérré (2003).

Records: M01 (2 males, 4 females); M06 (1 male); M12 (2 males, 1 female); M55 (1 female). N = 11 

adults; 5 males, 6 females.

Methods: litter condenser-Berlese funnel (1 female); pitfall traps (4 males, 3 females); search (1 male, 

2 females).

Habitats: litter, freshwater marsh edge (2 males, 1 female); litter, seashore (1 male, 1 female); litter, 

deciduous-coniferous woodland (1 female); litter, saltmarsh (1 male); sifted grass litter near freshwater 

stream (1 female); running on soil in vegetable garden (1 female); in shed (1 female); on wall, house 

(1 male).

Regional Distribution: ME (Emerton 1909, as Micaria gentiles Banks); NH, CT, NY, ON, QC, NS 

(Platnick and Shadab 1988); NB, NF (Platnick and Dondale 1992).

Sergiolus ocellatus (Walckenaer, 1837)

Taxonomy: Platnick and Shadab (1981); Platnick and Dondale (1992); Paquin and Dupérré (2003).

Records: M06 (7 males, 3 females); M07 (1 male); M12 (1 male); M26 (1 male, 2 females); M35 

(1 male). N = 16 adults; 11 males, 5 females.

Methods: litter condenser-Berlese funnel (1 male); pitfall traps (9 males, 5 females); sweep net 

(1 male).

Months: June (2 males); July (9 males, 2 females); July-August (2 females); August (1 female).

Habitats: litter, saltmarsh (7 males, 3 females); litter, saltmarsh edge (1 male, 2 females); litter, 

Kalmia-Vaccinium heath (1 male); sifted marsh litter (grasses, rushes), brackish marsh (1 male); swept 

from marsh grasses and shrubs (Spiraea-Kalmia), freshwater marsh (1 male).

Regional Distribution: ME (Sferra et al., unpubl.); MA, NY, ON, NB, NS (Platnick and Shadab 

1981).

Zelotes exiguoides Platnick & Shadab, 1983

Taxonomy: Platnick and Shadab (1983); Platnick and Dondale (1992).

Records: M01 (2 females); M07 (6 males, 7 females). N = 15 adults; 6 males, 9 females.

Method: pitfall traps (6 males, 9 females).
125

Months: June (2 males, 4 females); June-July (2 females); July (4 males, 3 females).

Habitats: litter, Kalmia-Vaccinium heath (6 males, 7 females); litter, Betula-Acer stand (1 female); 

litter, Amelanchier-Rubus stand (1 female).

Regional Distribution: ME (Sferra et al., unpubl.); NH, ON (Platnick and Shadab 1983).

Zelotes fratris Chamberlin, 1920

Taxonomy: Platnick and Shadab (1983); Platnick and Dondale (1992); Paquin and Dupérré (2003).

Records: M01 (11 males, 10 females); M04 (2 females); M05 (1 female); M06 (22 males, 22 

females); M07 (10 males, 4 females); M12 (5 males, 3 females); M15 (1 female); M19 (1 female); 

M22 (11 males, 2 females); M24 (4 males, 1 female); M25 (8 males, 8 females); M26 (12 males, 18 

females); M27 (7 males, 5 females); M42 (2 males, 1 female); M56 (1 female). N = 172 adults; 92 

males, 80 females.

Methods: litter condenser-Berlese funnel (1 female); litter condenser-hand sorted (1 female); pitfall 

traps (82 males, 69 females); search (9 males, 8 females); unsifted litter-Berlese funnel (1 male); n. d. 

(1 female).

Months: May (12 males, 3 females); May-June (2 males, 9 females); June (22 males, 13 females); 

June-July (3 males, 2 females); July (22 males, 24 females); July-August (7 males, 7 females); August 

(19 males, 18 females); August-September (5 males, 1 female); September (3 females).

Habitats: litter, saltmarsh (22 males, 22 females); litter, saltmarsh edge (12 males, 18 females); 

litter, white (paper) birch stand (8 males, 8 females); litter, Kalmia-Vaccinium heath (9 males, 4 

females); litter, bigtooth aspen stand (11 males, 2 females); litter, red maple-white (paper) birch stand 

(7 males, 5 females); litter, freshwater marsh edge (5 males, 3 females); litter, red maple sapling stand 

(4 males, 1 female); litter, Betula-Acer stand (3 males, 1 female); litter, edge of old fi eld and alder 

swamp (1 male, 3 females); litter, seashore-backshore (2 females); unsifted litter, Kalmia-Vaccinium 

heath (1 male); sifted litter, saltmarsh wrack line (1 female); sifted litter-hand sorted, salt meadow 

(1 female); under rocks near park drive, coastal spruce-mixed hardwood (2 females); under rock in 

silk retreat, open ledges in mixed conifer, island (1 female); under rock on ledges, red spruce-mixed 

conifer, island (1 male); on ledge under beating cloth, mixed conifer, island (1 male); old fi eld 

(1 female); running across gravel lane (1 female); “dead” carried by ant, road (1 female); in bathtub, 

house (4 males, 1 female); in garage, house (1 male); on cellar door, house (1 female); on fl oor, house 

(1 male); in sun room, house (1 male); under tarps in screen house (1 female).

Regional Distribution: ME (Bishop 1923, as Zelotes ater (Hentz)); MA, CT, NH, VT, NY, ON, 

QC, NB, NS, PE (Platnick and Shadab 1983); NF (Platnick and Dondale 1992).

Zelotes hentzi Barrows, 1945

Taxonomy: Platnick and Shadab (1983); Platnick and Dondale (1992); Paquin and Dupérré (2003).

Records: M06 (4 females); M07 (2 males, 2 females); M10 (1 female); M26 (6 males, 1 female). N = 

16 adults; 8 males, 8 females.
126

Methods: pitfall traps (8 males, 7 females); search (1 female).

Months: May (3 males, 2 females); May-June (1 male); June (3 males, 2 females); July (1 male, 

4 females).

Habitats: litter, saltmarsh edge (6 males, 1 female); litter, saltmarsh (4 females); litter, KalmiaVaccinium heath (2 males, 2 females); under rocks on ledge; mixed conifer-hardwood (1 female).

Regional Distribution: ME (Collins et al. 1996); MA, CT, NH, NY, ON, NB, NS (Platnick and 

Shadab 1983); QC (Platnick and Dondale 1992).

Note: Emerton’s (1890) record of Prosthesima atra (sic.) from Washington County, ME may refer to 

either Zelotes fratris Chamberlin, 1920 (males) or Zelotes hentzi Barrows, 1945 (females), or to both 

species; see Kaston’s (1981) comments under Zelotes subterraneus (C. L. Koch, 1839), a European 

species. Likewise, Bryant’s (1908) records of Melanophora atra (sic.) (Hentz) Simon from Lincoln, 

Piscataquis, and Washington Counties may refer to either Z. fratris or Z. hentzi, or to both species.
127

FAMILY PHILODROMIDAE

The philodromids or running crab spiders are swift runners over and among vegetation and other 

surfaces where they hunt their prey by active pursuit or ambush. They typically have fl attened 

bodies, and their laterigrade legs are nearly equal in length and thickness. The legs are equipped with 

claw tufts and scopulae that allow quick, erratic movements on slippery and precipitous surfaces of 

plants, rocks, and ledges. The females spin silken egg sacs among needles of conifers or in folds of 

hardwood leaves; the females generally remain with their egg sacs until the spiderlings emerge. Most 

philodromids overwinter in the penultimate stage, sometimes in loose aggregations or “pseudofl ocks.” 

They generally reach maturity in early spring.

Formerly considered as the subfamily Philodrominae of the family Thomisidae, the philodromids are 

easily distinguished by their somewhat elongate fl attened bodies; long slender scopulate legs, with legs 

I, III, and IV nearly equal in length; eyes usually uniform in size and without prominent eye tubercles 

and arranged in two recurved rows, with the posterior row more recurved than the anterior; body 

with soft, recumbent setae. Additional defi ning characters, notes on natural and taxonomic histories, 

and a key to the genera of North America are given by Dondale (2005b).

C. D. Dondale and J. H. Redner, formerly of the Biosystematics Research Institute, Ottawa, revised 

most of the genera and species of Philodromidae found in North America. Sauer and Platnick 

(1972) revised the genus Ebo. These revisions were published during 1961-1976; see Dondale and 

Redner 1978b for a listing of citations. Dondale and Redner (1978b) also provide identifi cation 

keys, descriptions, and illustrations of the genera and species found, or likely to be found, in the 

northeastern region. The philodromid fauna in Maine consists of 4 genera and 20 species; 3 genera 

and 13 species are found in Milbridge. Thus far, species of Ebo remain to be found in Milbridge; E. 

iviei Sauer & Platnick 1972 has been taken in nearby Jonesboro (Collins et al. 1996).

Philodromus cespitum (Walckenaer, 1802)

Taxonomy: Dondale (1961); Dondale and Redner (1976a); Dondale and Redner (1978b); Paquin 

and Dupérré (2003).

Records: M01 (1 female); M51 (2 females). N = 3 adult females.

Methods: beating cloth (1 female); search (1 female); shake-sweep net (1 female).

Months: June (2 females); July (1 female).

Habitats: on poplar leaf, seashore (1 female); shaking speckled alder bushes, mixed conifer opening 

(1 female); beating eastern larch foliage, mixed conifer (1 female).

Regional Distribution: ME (Dondale 1961, as Philodromus cespiticolis Walckenaer); NH, VT, MA, 

NY, ON, QC, NB, NS (Dondale and Redner 1976a).

Philodromus exilis Banks, 1892

Taxonomy: Dondale and Redner (1968); Dondale and Redner (1978b); Paquin and Dupérré (2003).
128

Records: M01 (2 males, 1 female); M06 (1 male, 1 female); M10 (1 male); M23 (2 females); M38 (1 

female); M51 (1 female); M55 (1 male). N = 11 adults; 5 males, 6 females.

Methods: beating cloth (5 males, 4 females); search (1 female); sweep net (1 female).

Months: May (4 males); June (1 male, 1 female); July (3 females); August (2 females).

Habitats: beating red spruce foliage, mixed conifer, island (2 females); beating red spruce saplings, 

conifer-mixed hardwood (1 male); beating red spruce, near stream (1 male); beating white spruce 

foliage, edge of saltmarsh (1 male, 1 female); beating spruces, old fi eld edge (2 males); beating 

eastern larch, mixed conifer (1 female); sweeping Sphagnum bog (1 female); on rolled, dead alder leaf 

(1 female).

Regional Distribution: ME, NH, VT, NY, ON, QC, NB, NS (Dondale and Redner 1968).

Philodromus imbecillus Keyserling, 1880

Taxonomy: Dondale and Redner (1968, 1976a); Paquin and Dupérré (2003).

Records: M06 (1 female); M28 (1 male); M35 (1 male). N = 3 adults; 2 males, 1 female.

Method: sweep net (2 males, 1 female).

Months: June (1 male, 1 female); July (1 male).

Habitats: sweeping herbaceous vegetation, saltmarsh (1 female); sweeping roadside disturbed area 

(1 male); sweeping grasses and rushes, brackish marsh (1 male).

Regional Distribution: ME (Bryant 1908, as Philodromus lineatus Emerton, 1892); NH, VT, MA, 

CT, NY, ON, QC, NS, LB (Dondale and Redner 1968).

*Philodromus peninsulanus Gertsch, 1934

Taxonomy: Dondale and Redner (1968, 1976a); Paquin and Dupérré (2003).

Records: M12 (6 females); M33 (3 males, 4 females); M55 (1 female). N = 14 adults; 3 males, 11 

females.

Method: sweep net (3 males, 11 females).

Months: June (2 males, 2 females); July (1 male, 8 females); August (1 female).

Habitats: sweeping grasses and shrubs (Spiraea sp.), freshwater marsh (6 females); sweeping grasses 

and sedges, freshwater marsh (3 males, 4 females); sweeping grasses, streamside (1 female).

Regional Distribution: ME (this study); NY, ON (Dondale and Redner 1968, 1978b); QC 

(Bélanger and Hutchinson 1992).
129

Philodromus placidus Banks, 1892

Taxonomy: Dondale and Redner (1968, 1976a); Paquin and Dupérré (2003).

Records: M01 (1 male, 3 females); M02 (1 female); M49 (1 male); M51 (1 female). N = 7 adults; 

2 males, 5 females.

Methods: beating cloth (1 male, 5 females); sweep net (1 male).

Months: June (1 male, 4 females); July (1 male); August (1 female).

Habitats: beating red spruce foliage (3 females); beating red spruce, seashore-backshore (1 male); 

beating red spruce, mixed conifer (1 female); beating young balsam fi r foliage, mixed conifer 

(1 female); sweeping old fi eld (1 male).

Regional Distribution: ME (Bryant 1908, as Philodromus bidentatus Emerton); NH, VT, MA, CT, 

ON, QC, NB, NS, LB (Dondale and Redner 1968).

Philodromus praelustris Keyserling, 1880

Taxonomy: Dondale (1961); Dondale and Redner (1976a); Paquin and Dupérré (2003).

Records: M01 (1 male, 1 female). N = 2 adults; 1 male, 1 female.

Method: search (1 male, 1 female).

Months: May (1 male); June (1 female).

Habitats: outside wall of house (1 female); in shed (1 male).

Regional Distribution: ME (Procter 1946); NH, MA, CT, NY, ON, QC, NB, NS (Dondale and 

Redner 1976a).

Philodromus rufus vibrans Dondale, 1964

Taxonomy: Dondale and Redner (1968, 1976a); Paquin and Dupérré (2003).

Records: M01 (16 males, 14 females); M10 (1 female); M12 (1 female); M23 (1 female); M35 

(2 females); M50 (1 female); M51 (2 females). N = 38 adults; 16 males, 22 females.

Methods: beating cloth (8 males, 6 females); search (6 males, 3 females); sweep net (2 males, 13 

females). 

Months: March (1 male); May (13 males, 6 females); June (2 males, 7 females); July (7 females); 

August (2 females).

Habitats: beating red spruce foliage, old fi eld edge (1 male, 5 females); beating red spruce, conifermixed hardwood (1 female); beating spruces, old fi eld edge (7 males); sweeping old fi eld, inland 
130

(2 males, 5 females); sweeping old fi eld (Kalmia sp.), abandoned farm on island (1 female); sweeping 

low vegetation, deciduous-coniferous woodland (1 female); sweeping grasses and shrubs, freshwater 

marsh (1 female); sweeping brackish marsh (2 females); sweeping grasses, forbs, and shrubs, seashorebackshore (1 female); sweeping meadowsweet, mixed conifer opening (2 females); on bedroom ceiling 

(1 female), terrace (1 female); boat (1 female); house walls (2 males); offi ce desk (1 male), lawn chair 

(1 male); in woodpile (1 male), old fi eld (1 male).

Regional Distribution: ME (Bryant 1908); NH, VT, MA, CT, RI, ON, QC, NB, NS (Dondale and 

Redner 1968).

Note: Bryant (1908) recorded Philodromus pictus Emerton from Maine; however, Dondale and 

Redner (1976a) list Emerton’s P. pictus as a synonym of Philodromus rufus vibrans Dondale, 1964.

Philodromus vulgaris (Hentz, 1847)

Taxonomy: Dondale (1961); Dondale and Redner (1976a); Paquin and Dupérré (2003).

Record: M01 (1 female). N = 1 adult female.

Method: search (1 female).

Month: May (1 female).

Habitat: inside screen door (1 female).

Regional Distribution: ME (Bryant 1908, as Philodromus vulgaris (Hentz) Keyserling); MA, CT, RI, 

NY, ON, QC, NS (Dondale 1961).

Thanatus formicinus (Clerck, 1757)

Taxonomy: Dondale et al. (1964); Dondale and Redner (1978b); Paquin and Dupérré (2003).

Records: M01 (1 male); M05 (1 female); M06 (1 male); M07 (1 male, 2 females); M26 (2 females). 

N = 8 adults; 3 males, 5 females.

Methods: pitfall traps (3 males, 4 females); sweep net (1 female).

Months: May (1 male); June (2 females); July (2 females); August (1 female); September (2 males).

Habitats: Kalmia-Vaccinium heath, litter (1 male, 2 females); saltmarsh edge, litter (2 females); 

saltmarsh, litter (1 male); old fi eld-alder swamp, litter (1 male); sweeping old fi eld (1 female).

Regional Distribution: ME (Procter 1938, as Thanatus formicinus (Oliver) (lycosoides Emerton; not 

coloradensis Keyserling); NH, VT, MA, CT, NY, ON, NS, NF (Dondale et al. 1964); QC (Bélanger 

and Hutchinson 1992).
131

Thanatus rubicellus Mello-Leitão, 1929

Taxonomy: Dondale et al. (1964); Dondale and Redner (1978b); Paquin and Dupérré (2003).

Records: M06 (2 males); M07 (2 males). N = 4 adult males.

Method: pitfall traps (4 males).

Months: May (1 male); July (3 males).

Habitats: Kalmia-Vaccinium heath, litter (2 males); saltmarsh, litter (2 males).

Regional Distribution: ME (Bryant 1908, as Thanatus rubicundus Keyserling); NH, VT, MA, CT, 

NY, QC, LB (Dondale et al. 1964).

Thanatus striatus C. L. Koch, 1845

Taxonomy: Dondale et al. (1964); Dondale and Redner (1978b); Paquin and Dupérré (2003).

Records: M06 (2 females); M26 (1 female); M35 (1 female). N = 4 adult females.

Methods: pitfall traps (2 females); sweep net (2 females).

Months: May (1 female); June (2 females); July (1 female).

Habitats: saltmarsh, litter (2 females); sweeping saltmarsh edge (1 female); sweeping grasses and 

rushes, brackish marsh (1 female).

Regional Distribution: ME, NH, MA, CT, ON, NS (Dondale et al. 1964); QC (Bélanger and 

Hutchinson 1992).

Tibellus maritimus (Menge, 1875)

Taxonomy: Dondale and Redner (1978b); Kaston (1981); Paquin and Dupérré (2003).

Records: M04 (1 male); M05 (4 males, 3 females); M06 (1 male, 4 females); M12 (13 males, 

7 females); M26 (2 males); M33 (2 males, 3 females); M35 (1 male); M47 (2 females); M55 

(2 males). N = 45 adults; 26 males, 19 females.

Method: sweep net (26 males, 19 females).

Months: May (4 males, 2 females); June (21 males, 12 females); July (1 male, 2 females); August 

(3 females).

Habitats: sweeping grasses, sedges, and shrubs (Spiraea, Kalmia), freshwater marshes (15 males, 

10 females); sweeping saltmarsh (Juncus, Spartina) (1 male, 5 females); sweeping saltmarsh edge 

(2 males); sweeping old fi eld grasses and forbs (4 males, 1 female); sweeping low wet area, coastal 

spruce-mixed hardwood (1 male); sweeping wet meadow (1 female); sweeping grasses and forbs, 
132

playground edge (2 males); sweeping vegetation, streamside (2 females); sweeping grasses and rushes, 

brackish marsh (1 male).

Regional Distribution: ME (Mairs and Jennings, unpubl.); CT (Kaston 1981); New England, ON, 

QC, NB, NS, NF, LB (Dondale and Redner 1978b).

Tibellus oblongus (Walckenaer, 1802)

Taxonomy: Dondale and Redner (1978b); Kaston (1981); Paquin and Dupérré (2003).

Records: M01 (10 males, 19 females); M06 (1 male, 5 females); M12 (1 male); M14 (1 male, 

1 female); M17 (1 male); M23 (2 males, 6 females); M28 (2 females); M35 (1 male, 2 females); M55 

(1 male). N = 53 adults; 18 males; 35 females.

Methods: search (6 males, 7 females); sweep net (12 males, 27 females); n. d. (1 female).

Months: May (2 males); June (5 males, 7 females); July (11 males, 17 females); August (11 females).

Habitats: sweeping old fi eld (3 males, 11 females); sweeping saltmarsh (1 male, 4 females); sweeping 

grasses and rushes, brackish marsh (1 male, 2 females); sweeping freshwater pond edge (2 males, 

1 female); sweeping blue-joint grass, seashore-backshore, island (3 females); sweeping sea oats, 

seashore-backshore, island (1 female); sweeping white and yellow clover, roadside disturbed area 

(2 females); sweeping mixed vegetation, school playground (1 male); sweeping bog near seashore, 

island (1 male, 1 female); sweeping grasses and forbs, mixed hardwood-conifer (1 male, 1 female); 

sweeping grasses and Kalmia sp., mixed conifer opening, island (1 male); sweeping grasses and 

meadowsweet, freshwater marsh (1 male); sweeping grass, island (1 female); on goldenrod, old fi eld (1 

female); on aster leaf, old fi eld (1 female); on astilbe leaf, garden (1 female); on iris leaf (1 female); on 

lettuce, garden (1 female); running on ground, garden (1 male); in compost pile, garden (1 female); 

in crevice of boat trailer parked in old fi eld (1 female); in old fi eld (1 male); on backyard rock 

(1 male); on screen door, house (1 male); inside kitchen window, house (1 male); in grass on beach, 

seashore (1 male); saltmarsh (1 female).

Regional Distribution: ME (Bryant 1908); CT (Kaston 1981); ON, QC, NB, NS (Dondale and 

Redner 1978b).
133

FAMILY THOMISIDAE

The thomisids or crab spiders have stout, moderately fl attened bodies, and strong laterigrade legs, 

especially legs I and II. Their movements are slow, deliberate, and crablike. They live in diverse 

habitats, including foliage of grasses, forbs, shrubs, and trees, and the litter of old fi elds, marshes, 

bogs, and forests. Some species are arboreal, inhabiting the mid- and upper-crowns of trees; others 

are strictly ground dwellers, or inhabitants of bark furrows and crevices. Brightly colored species of 

Misumena, Misumenoides, and Misumenops are frequently associated with fl ower blossoms where they 

capture prey by deception and ambush.

The thomisids can be distinguished from other crablike spiders by their stout bodies that are 

sparsely covered with simple, erect setae; their legs are devoid of scopulae and claw tufts; legs I and 

II are longer and more laterigrade than legs III and IV; and the lateral eyes are seated on prominent 

tubercles. Additional family character descriptors, notes on natural and taxonomic histories, and a key 

to the genera in North America are given by Dondale (2005c).

Species descriptions and generic revisions of North American Thomisidae are provided by Gertsch 

(1939, 1953), Schick (1965), and Turnbull et al. (1965). Dondale and Redner (1978b) provide 

identifi cation keys, descriptions, and illustrations of thomisid taxa likely to be found in the 

northeastern region. The Maine thomisid fauna consists of 8 genera and 29 species; 6 genera and 19 

species are found in Milbridge, including a new, undescribed or invasive species of Ozyptila. 

Coriarachne utahensis (Gertsch, 1932)

Taxonomy: Gertsch (1939); Schick (1965); Bowling and Sauer (1975); Dondale and Redner 

(1978b); Kaston (1981); Paquin and Dupérré (2003, as Bassaniana utahensis (Gertsch, 1932)); 

Dondale (2005c).

Records: M01 (5 males, 4 females); M05 (1 male); M21 (1 male); M35 (1 male). N = 12 adults; 

8 males, 4 females.

Methods: beating cloth (1 female); search (6 males, 2 females); sweep net (2 males, 1 female).

Months: April (1 female); May (2 males); June (3 males); July (2 males); October (1 male, 2 females); 

November (1 female).

Habitats: beating spruces (1 female); sweeping grasses and forbs in open fi eld, old fi eld (1 male); 

sweeping spruce foliage (1 female); sweeping vegetation, brackish marsh (1 male); in basement, 

house (1 male); in bathroom basin, house (1 male); in kitchen sink, house (1 female); in automobile 

(1 male); on bedroom ceiling, house (1 male); on wall, offi ce (1 male); on cellar wall, house 

(1 female); on exterior white vinyl siding, building (1 male).

Regional Distribution: ME (Gertsch 1953); NH, VT, MA, NY, ON, QC, NB, NS, LB (Bowling 

and Sauer 1975).

Notes: Ono (1988) assigned three of the four North American species of Coriarachne to the genus 

Bassaniana, including Coriarachne utahensis (Gertsch 1932). However, Dondale (2005c) indicated 
134

that numerous investigators have addressed this division and all agree that the four North American 

species are congeneric. The authors agree with Dondale’s assessment and prefer the more familiar 

name Coriarachne utahensis (Gertsch, 1932).

Misumena vatia (Clerck, 1757)

Taxonomy: Gertsch (1939, as Misumena calycina (Linnaeus, 1758); Schick (1965); Kaston (1981); 

Dondale and Redner (1978b); Paquin and Dupérré (2003).

Records: M01 (13 males, 12 females); M04 (1 male, 2 females) M06 (1 male, 1 female); M12 (2 

males); M28 (2 males, 1 female); M35 (1 male, 3 females); M51 (1 male); M55 (1 male). N = 41 

adults; 22 males, 19 females.

Methods: beating cloth (1 male, 1 female); search (8 males, 8 females); sweep net (13 males, 10 

females).

Months: May (1 male); June (10 males, 5 females); July (9 males, 10 females); August (2 males, 2 

females); September (2 females).

Habitats: beating spruces, old fi eld edge (1 male); beating spruce, saltmarsh edge (1 female); 

sweeping vegetation, old fi eld (3 males, 2 females); sweeping vegetation, old fi eld edge (1 male, 

1 female); sweeping vegetation, freshwater pond edge (1 female); sweeping roadside grasses, forbs, 

and shrubs, red spruce forest (1 male, 1 female); sweeping shrubby vegetation, coastal spruce-mixed 

hardwood (1 female); sweeping vegetation (Juncus-Spartina), saltmarsh (1 male); sweeping grasses and 

meadowsweet, freshwater marsh (1 male); sweeping grasses and shrubs, freshwater marsh (1 male); 

sweeping white and yellow clover, roadside disturbed area (2 males, 1 female); sweeping marsh 

vegetation, brackish marsh (1 male, 3 females); sweeping rhodora and meadowsweet, mixed conifer 

opening (1 male); sweeping grasses and forbs, school playground edge, disturbed area (1 male); in 

low vegetation, freshwater pond edge (1 female); in folded quaking aspen leaf (1 female); in and 

on goldenrod, old fi eld (1 male, 1 female); in rugosa rose fl ower (1 female); on aster leaves along 

lane, mixed hardwood-conifer (1 male, 1 female); dead, among grasses on incoming tide, seashore 

(1 female); on alder at shore, seashore-backshore (1 male); on marsh grasses, seashore (1 female); on 

raspberry cane, old fi eld (1 male); on ox-eye daisy, old fi eld (1 male); on ornamental daisy, fl ower 

garden (1 male); on foliage, fl ower garden (1 male); in kitchen, house (1 female); running on house 

deck, with prey (1 male).

Regional Distribution: ME (Packard 1905, as Misumena vatia Thorell); NH, VT, MA, NY, ON, 

QC, NS (Gertsch 1939, as Misumena calycina (Linnaeus)); CT (Kaston 1981); NB, NF, LB (Dondale 

and Redner 1978b).

Notes: The biology of Misumena vatia has been studied extensively by D. H. Morse and associates 

in Lincoln County, Maine. For example, see Morse (1985, 1986, 1991, 1992) and Morse and Fritz 

(1982).

Misumenops asperatus (Hentz, 1847)

Taxonomy: Gertsch (1939); Schick (1965); Dondale and Redner (1978b); Kaston (1981); Paquin 

and Dupérré (2003).
135

Record: M06 (1 female). N = 1 adult female.

Method: sweep net (1 female).

Month: July (1 female).

Habitat: sweeping vegetation, saltmarsh edge (1 female).

Regional Distribution: ME (Blake 1927, as Misumena asperata Emerton); NH, MA, CT, RI, NY, 

ON, QC (Gertsch 1939). 

*Misumenops carletonicus Dondale & Redner, 1976

Taxonomy: Dondale and Redner (1976b, 1978b); Paquin and Dupérré (2003).

Records: M01 (1 male); M09 (1 female). N = 2 adults; 1 male, 1 female.

Method: sweep net (1 male, 1 female).

Months: May (1 male); July (1 female).

Habitats: sweeping vegetation, old fi eld (1 male); sweeping vegetation, including sheep laurel, old 

fi eld-abandoned farm, island (1 female).

Regional Distribution: ME (this study); NY, ON (Dondale and Redner 1978b); QC (Hutchinson 

1999).

Ozyptila distans Dondale & Redner, 1975

Taxonomy: Gertsch (1939, as Ozyptila americana Banks); Dondale and Redner (1975, 1978b); 

Paquin and Dupérré (2003).

Records: M01 (4 males); M07 (6 males); M12 (1 male, 1 female); M22 (1 male, 2 females); M24 

(9 males, 2 females); M25 (9 males, 4 females); M26 (1 male); M27 (3 males). N = 43 adults; 34 

males, 9 females.

Method: pitfall traps (34 males, 9 females).

Months: May (3 males); May-June (6 males, 1 female); June (20 males, 5 females); June-July 

(5 males, 1 female); August (2 females).

Habitats: litter, seashore-backshore (1 male); litter, deciduous-coniferous woodland (3 males); litter, 

Kalmia-Vaccinium heath (6 males); litter, freshwater marsh edge (1 male, 1 female); forest-fl oor 

litter, bigtooth aspen stand (1 male, 2 females); forest-fl oor litter, red maple sapling stand (9 males, 

2 females); forest-fl oor litter, white (paper) birch stand (9 males, 4 females); litter, saltmarsh edge 

(1 male); forest-fl oor litter, red maple-white birch stand (3 males).
136

Regional Distribution: ME (Gertsch 1939, as Ozyptila americana Banks); NH, MA, CT, NY, ON, 

QC, NB, NS, PE, NF (Dondale and Redner 1978b).

**Ozyptila sp. (near distans Dondale & Redner, 1975)

Taxonomy: New or introduced species?

Record: M26 (1 male). N = 1 adult male.

Method: pitfall trap (1 male).

Month: June (1 male).

Habitat: litter, saltmarsh edge (1 male).

Regional Distribution: ME (this study); otherwise, unknown.

Note: This male specimen was examined by C. D. Dondale who concluded that it might be a new 

species or possibly an introduced species not previously found in North America (pers. comm.). It 

belongs to the brevipes group as defi ned by Dondale and Redner (1975). Like O. distans, the retrolateral 

tibial apophysis is not bent ventrad and does not extend beyond the mid-length of the tegulum. Distally, 

the ventral tibial apophysis is knobbed-shaped. The embolus is more angular than that of O. distans. The 

basal tegular ridge is less broad and bears a rounded tooth distally. Additional specimens, including those 

of both sexes, are needed for comparison with the types of Ozyptila distans Dondale & Redner, 1975, 

and with related species. The specimen from Milbridge is in the Canadian National Collection, Ottawa.

Tmarus angulatus (Walckenaer, 1837)

Taxonomy: Gertsch (1939); Dondale and Redner (1978b); Kaston (1981); Paquin and Dupérré 

(2003).

Records: M01 (2 males, 4 females); M12 (1 female); M35 (1 male). N = 8 adults; 3 males, 5 females.

Methods: beating cloth (1 female); search (1 male, 1 female); sweep net (2 males, 3 females).

Months: May (1 male); June (2 males, 3 females); July (1 female); August (1 female).

Habitats: beating hardwoods, mixed hardwood-conifer (1 female); sweeping vegetation, freshwater 

pond edge (1 female); sweeping vegetation, old fi eld (1 male, 1 female); sweeping shrubs, freshwater 

marsh (1 female); sweeping grasses, brackish marsh (1 male); on top of post (1 female); on exterior 

wall, house (1 male).

Regional Distribution: ME (Bryant 1908, as Tmarus caudatus (Hentz) Keyserling); NH, VT, MA, 

CT, NY (Gertsch 1939); ON, QC, NB, NS (Dondale and Redner 1978b).

Xysticus alboniger Turnbull et al., 1965

Taxonomy: Gertsch (1939, as Synema alboniger Keyserling, 1884); Turnbull et al. (1965); Dondale 

and Redner (1978b); Paquin and Dupérré (2003).
137

Records: M01 (2 males, 4 females); M07 (5 males); M26 (1 male). N = 12 adults; 8 males, 4 females.

Methods: pitfall traps (5 males); search (1 male, 1 female); sweep net (2 males, 3 females).

Months: May (2 males); May-June (1 male); June (5 males, 4 females).

Habitats: litter, Kalmia-Vaccinium heath (5 males); sweeping vegetation, old fi eld (2 females); 

sweeping grasses, forbs, and shrubs, old fi eld (1 male, 1 female); sweeping vegetation, saltmarsh edge 

(1 male); in vegetable garden (1 female); on foundation, house (1 male).

Regional Distribution: ME (Collins et al. 1996); NH, MA, CT, NY (Gertsch 1939, as Synema 

bicolor Keyserling); ON (Turnbull et al. 1965).

Xysticus ampullatus Turnbull et al., 1965

Taxonomy: Turnbull et al. (1965); Dondale and Redner (1978b); Paquin and Dupérré (2003).

Record: M07 (1 male). N = 1 adult male.

Method: pitfall trap (1 male).

Month: July (1 male).

Habitat: litter, Kalmia-Vaccinium heath (1 male).

Regional Distribution: ME (Collins et al. 1996); ON, NS, NB (Dondale and Redner 1978b); QC 

(Bélanger and Hutchinson 1992).

Xysticus chippewa Gertsch, 1953

Taxonomy: Gertsch (1953); Turnbull et al. (1965); Dondale and Redner (1978b); Paquin and 

Dupérré (2003).

Records: M12 (5 males, 9 females); M33 (4 males); M55 (2 females). N = 20 adults, 9 males, 11 

females.

Methods: search (6 females); sweep net (9 males, 5 females).

Months: June (8 males, 5 females); July (1 male, 2 females); August (4 females).

Habitats: sweeping grasses, freshwater marsh (1 male); sweeping grasses and meadowsweet, 

freshwater marsh (1 male); sweeping marsh grasses and shrubs (Spiraea, Kalmia), freshwater marsh (3 

males, 5 females); sweeping grasses and sedges, freshwater marsh (4 males); in retreats, blue-joint grass 

seedheads, freshwater marsh (4 females); in retreats, blue-joint grass seedheads, freshwater streamside 

(2 females).

Regional Distribution: ME (Longcore, unpubl.); ON (Gertsch 1953); QC (Bélanger and 

Hutchinson 1992).
138

Xysticus elegans Keyserling, 1880

Taxonomy: Gertsch (1939, 1953); Turnbull et al. (1965); Dondale and Redner (1978b); Paquin and 

Dupérré (2003).

Records: M01 (10 males, 6 females); M07 (2 males); M12 (1 female); M22 (13 males, 1 female); 

M24 (21 males); M25 (34 males, 2 females); M26 (1 male); M27 (16 males, 2 females). N = 109 

adults; 97 males, 12 females.

Methods: pitfall traps (95 males, 7 females); search (2 males, 1 female); sweep net (4 females).

Months: May (11 males); May-June (13 males, 1 female); June (42 males, 6 females); June-July (19 

males, 1 female); July (12 males, 2 females); September (1 female); October (1 female).

Habitats: forest-fl oor litter, Betula-Acer stand (1 female); litter, deciduous-coniferous woodland (5 

males, 1 female); litter, Amelanchier-Rubus stand (2 males); litter, seashore-backshore (1 male); litter, 

Kalmia-Vaccinium stand (2 males); forest-fl oor litter, bigtooth aspen stand (13 males, 1 female); 

forest-fl oor litter, red maple stand (21 males); forest-fl oor litter, white (paper) birch stand (34 males, 

2 females); litter, saltmarsh edge (1 male); forest-fl oor litter, red maple-white birch stand (16 males, 

2 females); sweeping vegetation, old fi eld (1 female); sweeping low understory shrubs, Betula-Acer 

woods (1 female); sweeping shore grasses and sedges, seashore-backshore (1 female); sweeping marsh 

grasses and shrubs (Spiraea-Kalmia), freshwater marsh (1 female); on exterior of garage door, house 

(1 female); on porch at night, house (1 male); on exterior wall, house (1 male).

Regional Distribution: ME (Bryant 1908, as Xysticus limbatus Keyserling); NH, VT, MA, CT, NY, 

ON, QC, NS (Gertsch 1939); NB, NF (Turnbull et al. 1965).

Note: Bryant’s (1908) record of Xysticus limbatus Keyserling from Cumberland County, ME may refer 

to X. elegans Keyserling (females only) or to X. emertoni Keyserling (males only), or to both species. 

Gertsch (1939) cited Bryant’s (1908) paper under both X. elegans and X. emertoni.

Xysticus ellipticus Turnbull et al., 1965

Taxonomy: Turnbull et al. (1965); Dondale and Redner (1978b); and Paquin and Dupérré (2003).

Records: M01 (1 male); M07 (3 males, 1 female); M26 (1 male). N = 6 adults; 5 males, 1 female.

Method: pitfall traps (5 males, 1 female).

Months: May (1 female); June (4 females); July (1 male).

Habitats: litter, Kalmia-Vaccinium heath (3 males, 1 female); litter, Amelanchier-Rubus stand 

(1 male); litter, saltmarsh edge (1 male).

Regional Distribution: ME (Jennings et al., unpubl.); NH, VT (Gertsch 1939, as Synema obscurum 

Keyserling); NB (Dondale and Redner 1978b); QC (Koponen 1994).
139

Xysticus emertoni Keyserling, 1880

Taxonomy: Gertsch (1939, 1953); Turnbull et al. (1965); Dondale and Redner (1978b); Kaston 

(1981); Paquin and Dupérré (2003).

Records: M01 (4 males, 17 females); M05 (2 females); M06 (1 female); M07 (2 females); M12 (2 

females); M20 (1 male); M37 (1 female); M55 (1 female). N = 31 adults; 5 males, 26 females.

Methods: pitfall trap (1 female); search (4 males, 7 females); sweep net (1 male, 17 females); n. d. 

(1 female).

Months: May (2 males, 2 females); June (2 males, 6 females); July (1 male, 8 females); August (9 

females); September (1 female).

Habitats: litter, Kalmia-Vacciniium heath (1 female); sweeping vegetation, old fi elds (1 male, 12 

females); sweeping vegetation, saltmarsh (1 female); sweeping grasses, freshwater marsh (1 female); 

sweeping marsh grasses and shrubs (Spiraea-Kalmia), freshwater marsh (1 female); sweeping grasses, 

forbs, and meadowsweet, mown fi eld (1 female); sweeping grasses and forbs, school playground 

edge (1 female); in common St. Johnswort, old fi eld (1 female); in old fi elds (2 females); in rolled 

chokecherry leaf (1 female); in camp near seashore (1 female); on blackberry leaf, old fi eld (1 female); 

on wild raspberry along lane (1 male); on wild cherry (1 male); on heath plants, Kalmia-Vaccinium 

heath (1 female); on foundation, house (1 male); on paved parking lot (1 male); under old board 

feeding on lycosid (1 female).

Regional Distribution: ME (Bryant 1908, as Xysticus limbatus Keyserling); NH, VT, MA, NY, ON, 

QC (Gertsch 1939); NB, NS, NF (Turnbull et al. 1965).

Notes: Regarding the identity of X. limbatus Keyserling listed by Bryant (1908), see notation under 

Xysticus elegans above. The spider wasp Dipogon sayi sayi Banks provisioned its nests with paralyzed 

female crab spiders, chiefl y X. emertoni, in strip-clearcut stands of spruce-fi r-mixed hardwood on the 

Moosehorn National Wildlife Refuge, near Calais, ME (Jennings and Parker 1987).

Xysticus ferox (Hentz, 1847)

Taxonomy: Gertsch (1939, 1953); Turnbull et al. (1965); Kaston (1981); Dondale and Redner 

(1978); and Paquin and Dupérré (2003).

Records: M01 (2 males); M06 (1 male); M07 (2 males); M25 (1 male, 1 female); M26 (1 male). N = 

8 adults; 7 males, 1 female.

Methods: pitfall traps (5 males, 1 female); search (2 males).

Months: May (3 males, 1 female); June (3 males); July (1 male).

Habitats: litter, Kalmia-Vaccinium heath (2 males); litter, saltmarsh (1 male); litter, white (paper) 

birch stand (1 male, 1 female); litter, saltmarsh edge (1 male); on screen door, house (1 male); 

running in vegetable garden (1 male).
140

Regional Distribution: ME (Bryant 1908, as Xysticus stomachosus Keyserling); NH, VT, MA, CT, 

RI, NY, ON, LB (Gertsch 1939); QC, NS (Turnbull et al. 1965); NB (Dondale and Redner 1978).

Xysticus luctans (C. L. Koch, 1845)

Taxonomy: Gertsch (1939, 1953); Turnbull et al. (1965); Kaston (1981); Dondale and Redner 

(1978); Paquin and Dupérré (2003).

Records: M01 (2 males); M07 (2 males, 1 female). N = 5 adults; 4 males, 1 female.

Methods: pitfall traps (2 males, 1 female); search (2 males).

Months: May (2 males, 1 female); May-June (1 male); June (1 male).

Habitats: litter, Kalmia-Vaccinium heath (2 males, 1 female); on cobble beach, seashore (1 male); 

running in old fi eld (1 male).

Regional Distribution: ME (Bryant 1908, as Xysticus quadrilineatus Keyserling); VT, NH, MA, CT, 

NY, QC, NS (Gertsch 1939); ON (Gertsch 1953). 

**Xysticus luctuosus (Blackwall, 1836)

Taxonomy: Gertsch (1939), as Xysticus lutulentus Gertsch, 1934; Locket and Millidge (1951); 

Turnbull et al. (1965); Dondale and Redner (1978b); Paquin and Dupérré (2003).

Records: M01 (1 male); M30 (1 male); M55 (1 male). N = 3 adult males.

Methods: search (1 male); sweep net (2 males).

Months: June (2 males); July (1 male).

Habitats: sweeping roadside grasses, forbs, ferns, and rushes, alder-aspen edge (1 male); sweeping 

grasses and forbs, school playground edge (1 male); on window screen, house (1 male).

Regional Distribution: ME (this study); ON, QC (Turnbull et al. 1965); NB (Dondale and Redner 

1978); LB (Gertsch 1939, as Xysticus lutulentus Gertsch, 1934).

Xysticus punctatus Keyserling 1880

Taxonomy: Gertsch (1939, 1953); Turnbull et al. (1965); Dondale and Redner (1978b); Kaston 

(1981); Paquin and Dupérré (2003).

Records: M01 (7 males, 8 females); M05 (2 males); M06 (1 female); M37 (1 female); M44 (1 male); 

M57 (1 female). N = 21 adults; 10 males, 11 females.

Methods: beating cloth (6 males, 2 females); pitfall trap (1 female); search (1 male, 4 females); sweep 

net (3 males, 4 females).
141

Habitats: beating red spruce foliage (4 males, 1 female); beating red spruce foliage, old fi eld edge 

(2 males); beating spruces, old fi eld edge (1 female); litter, saltmarsh (1 female); sweeping grasses and 

forbs, old fi eld (1 male); sweeping vegetation, old fi eld (1 male, 2 females); sweeping wet meadow, old 

fi eld-brackish marsh edge (1 male); sweeping and shaking chokecherry foliage (1 female); sweeping 

rushes, brackish marsh (1 female); in folded chokecherry leaves, old fi eld (2 females); in rolled pin 

cherry leaf (1 female); on raspberry canes, garden (1 female); on rock wall, cemetery (1 male).

Regional Distribution: ME (Bryant 1908, as Xysticus formosus Banks); NH, VT, MA, NY, ON, NS 

(Gertsch 1939); CT (Gertsch 1953); ON, NB (Turnbull et al. 1965); QC (Bélanger and Hutchinson 

1992).

Xysticus triguttatus Keyserling, 1880

Taxonomy: Gertsch (1939, 1953); Turnbull et al. (1965); Dondale and Redner (1978); Kaston 

(1981); Paquin and Dupérré (2003).

Records: M01 (8 males, 5 females); M07 (2 males, 2 females); M23 (1 male). N = 11 males, 7 females.

Methods: pitfall traps (2 males, 1 female); search (4 males, 1 female); sweep net (5 males, 5 females).

Months: May-June (1 male); June (6 males, 1 female); June-July (1 male); July (3 males, 2 females); 

August (3 females); September (1 female).

Habitats: litter, Kalmia-Vaccinium heath (2 males, 1 female); sweeping grasses, forbs, and shrubs, old 

fi eld (2 females); sweeping vegetation, old fi eld (3 males, 2 females); sweeping vegetation, old fi eld 

edge (1 male); sweeping vegetation, Kalmia-Vaccinium heath (1 female); sweeping grasses and Kalmia, 

mixed conifer opening, island (1 male); on ox-eye daisy, old fi eld (1 female); in bathtub, house 

(1 male); in shed (1 male); on exterior wall, house (2 males).

Regional Distribution: ME (Bryant 1908); NH, MA, CT, NY, NS (Gertsch 1939); ON (Gertsch 

1953); QC, NF (Turnbull et al. 1965); NB (Dondale and Redner 1978). 

Xysticus winnipegensis Turnbull et al., 1965

Taxonomy: Turnbull et al. (1965); Dondale and Redner (1978b); Redner and Dondale (1980); 

Paquin and Dupérré (2003).

Records: M07 (11 males). N = 11 adult males.

Method: pitfall traps (11 males).

Months: May (9 males); June (2 males).

Habitat: litter, Kalmia-Vaccinium heath (11 males).

Regional Distribution: ME (Collins et al. 1996); NB (Dondale and Redner 1978); QC (Bélanger 

and Hutchinson 1992).
142

FAMILY SALTICIDAE

The salticids or jumping spiders have the keenest eyesight of all spiders. They are diurnal hunters 

that slowly stalk their prey until a short distance away. They then make a sudden quick jump, with 

the front legs (usually the heaviest) extended and an anchoring line fastened to the substrate by the 

spinnerets. These strategies permit salticids to leap from vertical surfaces and capture fl ying insects. 

They are found in diverse habitats, including cobble beaches and rocks in old fi elds and meadows; 

on foliage of grasses, forbs, shrubs, and small trees; on and under bark of standing and dead trees, 

and bark of stumps; and on sandy beaches of streams, lakes, rivers, and seashores. They are usually 

common and active in open sunny places. Richman and Jackson (1992) reviewed the behavior of 

these most fascinating spiders.

None of our North American species of Salticidae are known to spin webs for prey capture. However, 

they all construct closely woven retreats for molting, hibernating, and egg-laying. Often several 

spiders of the same (or different) species will spin their hibernating nests in aggregations under bark; 

others spin maternal nests among dry leaves and on and in dry, senescent seedheads of forbs and 

shrubs.

The salticids are easily recognized by their large anterior median eyes and minute posterior median 

eyes. The eyes are typically arranged in three rows; the posterior row is strongly recurved, thus 

appearing as two rows. Legs I and II are prograde, some with enlarged femora; the tarsi are twoclawed, and claw tufts are usually present. Some species have constricted bodies and mimic ants 

(Cutler 1988); others have wide bodies and mimic beetles (Gertsch 1979, Richman and Jackson 

1992). Richman et al. (2005) provide additional information on family characteristics, natural and 

taxonomic histories, and a key to the genera in North America north of Mexico.

This is the largest family of spiders, with 560 genera and 5,077 species worldwide (Platnick 2007). 

Collectively, the United States and Canada have more than 315 species in 63 genera (Richman et 

al. 2005). The Maine salticid fauna is represented by 21 genera and 50 species; 15 genera and 25 

species are found in Milbridge. Source literature, including revisionary works, are listed below for 

each species found in Milbridge. Kaston (1981) provides an identifi cation key for the more common 

species likely to be found in New England; however, some of the species names are now out-of-date. 

See the latest version of Norman I. Platnick’s “World Spider Catalog” (e.g., Platnick 2007, version 

7.5) for currently approved names.

Eris militaris (Hentz, 1845)

Taxonomy: Maddison (1986); Kaston (1981, as Eris marginata (Walckenaer)); Paquin and Dupérré 

(2003).

Records: M01 (50 males, 64 females); M02 (1 male, 2 females); M04 (9 males, 5 females); M06 (8 

males, 7 females); M07 (3 males, 2 females); M10 (1 male); M12 (3 males, 7 females); M13 (1 male, 

1 female); M15 (2 females); M17 (1 female); M26 (1 female); M30 (2 males); M31 (1 female); M32 

(1 female); M33 (2 males, 3 females); M34 (1 male, 5 females); M35 (3 females); M37 (2 males, 5 

females); M38 (3 males, 2 females); M39 (2 males); M47 (1 male); M51 (2 males, 1 female); M55 

(1 male); M57 (1 female); M58 (1 female). N = 207 adults; 92 males, 115 females.
143

Methods: beating cloth (12 males, 15 females); brushing tree-bole bark (1 female); search (29 males, 

29 females); sifted litter-hand sorted (1 female); sweep net (49 males, 69 females); n. d. (2 males).

Months: May (1 male, 1 female); June (15 males, 23 females); July (9 males, 21 females); August 

(48 males, 35 females); September (18 males, 30 females); October (1 male, 4 females); December 

(1 female).

Habitats: beating red spruce foliage, seashore-backshore (2 males, 1 female); beating red spruce 

foliage, old fi eld edge (1 female); beating red spruce foliage, mixed conifer-hardwood (1 male); 

beating red spruce foliage, saltmarsh edge (2 females); beating red spruce foliage (1 male, 1 female); 

beating spruce branches (1 male); beating spruce at shore, seashore-backshore (2 males); beating 

spruces, saltmarsh edge (1 female); beating white pine foliage (1 male, 1 female); beating alders, 

old fi eld edge (1 female); beating foliage in alder thicket (2 males, 2 females); beating foliage of 

hardwoods (1 male, 3 females); beating eastern larch foliage (1 male, 2 females); brushing white 

(paper) birch bark, Betula-Acer stand (1 female); sifting forest-fl oor litter, Acer-Betula-Picea stand 

(1 female); sweeping grasses, forbs, shrubs, and small trees, old fi eld (5 males, 7 females); sweeping 

vegetation, old fi eld (15 males, 18 females); sweeping vegetation, old fi eld edge (1 male, 2 females); 

sweeping vegetation, freshwater pond shore (3 females); sweeping shore grasses and sedges, seashorebackshore (1 female); sweeping aster and raspberry (1 male); sweeping stand of meadowsweet 

(1 male); sweeping vegetation, low marshy area, coastal spruce-mixed hardwood (2 females); sweeping 

vegetation (Juncus-Spartina), saltmarsh (7 males, 7 females); sweeping vegetation, saltmarsh edge 

(1 female); sweeping vegetation, Kalmia-Vaccinium heath (2 males, 2 females); sweeping grasses, 

sedges, and shrubs, freshwater marsh (3 males, 7 females); sweeping grasses and sedges, freshwater 

marsh (1 male, 2 females); sweeping roadside grasses, forbs, ferns, and rushes, roadside alder-aspen 

edge (1 male); sweeping foliage, mixed hardwood-conifer (1 female); sweeping grasses, sedges, and 

forbs, dry streambed, mixed hardwood-conifer (1 male, 4 females); sweeping vegetation, brackish 

marsh (2 females); sweeping and shaking chokecherry foliage and fl owers, old fi eld (1 female); 

sweeping grasses and Spiraea, mown fi eld edge (2 males, 4 females); sweeping vegetation, Sphagnumcranberry-rush bog (3 males, 2 females); sweeping understory shrubs and ferns, red maple sapling 

stand (2 males); sweeping marsh vegetation near freshwater stream, riparian (1 male); sweeping 

meadowsweet, mixed conifer opening (2 males, 1 female); sweeping grasses and forbs, edge of school 

playground, disturbed area (1 male); sweeping rushes, brackish marsh (1 female); sweeping grasses, 

forbs, and rushes, brackish marsh (1 female); in old fi eld (1 male, 3 females); in grass (1 female); in 

retreat on chokecherry (1 female); in alders along shore, seashore-backshore (1 female); in silk nest, 

St. Johnswort fl ower head (1 male); in silk nest, folded blackberry leaf (1 male); in folded cherry 

leaf, old fi eld (1 male); in garden (1 male); in retreat with penultimate female, saltmarsh (1 male); in 

low foliage along streamside, riparian (1 male, 1 female); in folded red maple leaf, mixed hardwoodconifer (1 female); in silk retreat, folded green leaves, meadowsweet, freshwater marsh edge (1 

female); in silk retreat, dry meadowsweet infl orescence, freshwater marsh edge (1 male); in dead 

blackberry leaf (1 female); in downed burned trees over dry streambed, mixed hardwood-conifer 

(1 female); in nest, folded dead leaf impaled on black rush (Juncus sp.), brackish marsh (1 female); 

on grass blade, old fi eld edge (1 female); on vegetation, old fi eld (1 male); on potato plant, vegetable 

garden (1 female); on squash leaf, vegetable garden (1 female); on green bean leaf, vegetable garden (1 

male); on alder leaf and twig (2 females); on alder leaf along lane, mixed hardwood-conifer (1 male); 

on shrub foliage along lane, mixed hardwood-conifer (1 male); on foliage in alder thicket (1 male); on 

autumn olive (1 female); on mountain-ash feeding on bug (1 female); on blackberry leaves (1 female); 
144

on ground, old fi eld edge (1 male); on meadowsweet (1 male); on grass, freshwater pond edge 

(1 male); on rocks, seashore-backshore (2 males, 1 female); on balsam fi r foliage, understory balsam 

fi r-red spruce stand (1 male); descending on dragline in garage, house (1 female); on outside shingled 

wall, house (1 male, 2 females); on cellar door (exterior), house (1 female); on ceiling light fi xture 

(dead), house (1 female); on lawn chair feeding on winged ant (1 female); on lawn chair (1 male); 

on lawn table (1 male); on metal table, old fi eld (1 male); on table inside house (1 male); on porch, 

house (1 male); on screen door (1 male); running on brick wall (1 male); under bark, white (paper) 

birch, Betula-Acer stand (1 female); under bark, dead red maple, coastal spruce-mixed hardwood 

(1 male, 1 female); Kalmia-Vaccinium heath (1 male); n. d. (2 males).

Regional Distribution: ME (Bryant 1908, as Dendryphantes militaris (Hentz) Emerton); CT 

(Kaston 1981, as Eris marginata (Walckenaer)); NY (Crosby and Bishop 1928); QC (Bélanger and 

Hutchinson 1992).

Euophrys monadnock Emerton, 1891

Taxonomy: Kaston (1938b, 1981, as Euophrys nearctica Kaston, 1938); Edwards (1980); Paquin and 

Dupérré (2003).

Record: M01 (1 male). N = 1 adult male.

Method: search (1 male).

Month: June (1 male).

Habitat: on inside wall, house (1 male).

Regional Distribution: ME (Bryant 1908, as Evophrys (sic.) monadnock Emerton); NH (Richman 

and Cutler 1978, as Euophrys nearctica Kaston, 1938); QC (Paquin and Dupérré 2003).

Evarcha hoyi (Peckham & Peckham, 1883)

Taxonomy: Chickering (1944); Kaston (1981, as Evarcha hoyi Peckham)); Paquin and Dupérré 

(2003).

Records: M01 (22 males, 20 females); M03 (1 male); M05 (2 males, 2 females); M06 (5 males, 

4 females); M07 (1 male); M10 (2 females); M12 (1 female); M14 (2 males, 2 females); M15 (1 

female); M17 (1 female); M23 (3 females); M28 (2 females); M34 (1 female); M35 (6 females); M38 

(1 male); M39 (1 male); M41 (4 females); M51 (1 male, 1 female); M55 (3 males, 3 females); M57 

(1 female). N = 93 adults; 39 males, 54 females.

Methods: beating cloth (5 females); search (1 male, 14 females); sweep net (37 males, 34 females); n. 

d. (1 male, 1 female).

Months: May (3 females); June (6 males, 15 females); July (9 males, 14 females); August (11 males, 

11 females); September (13 males, 11 females).
145

Habitats: beating white pine foliage (2 females); beating red spruce foliage (saplings), conifer-mixed 

hardwood (2 females); beating wild-raisin, brackish marsh edge (1 female); sweeping grasses, forbs, 

shrubs, and small trees, old fi eld (12 males, 7 females); sweeping vegetation, old fi elds (7 males, 

7 females); sweeping Vaccinium and small shrubs, old fi eld (1 female); sweeping vegetation, old fi eld 

edge (2 males, 1 female); sweeping Vaccinium and Myrica near woods, old fi eld (1 male); sweeping 

low vegetation, deciduous-coniferous woodland understory (1 male); sweeping vegetation, saltmarsh 

(4 males, 2 females); sweeping grasses and shrubs, saltmarsh (1 female); sweeping vegetation, 

saltmarsh edge (1 female); sweeping grasses, saltmarsh edge (1 female); sweeping vegetation, KalmiaVaccinium heath (1 male); sweeping vegetation, freshwater marsh (1 female); sweeping understory 

vegetation along trail, mixed hardwood-conifer (2 males, 2 females); sweeping grass, island (2 

females); sweeping grassy area near cabin, mixed conifer opening, island (1 female); sweeping Kalmia 

near freshwater pond, island (1 male); sweeping roadside white and yellow clover, disturbed area 

(1 female); sweeping roadside disturbed area (1 female); sweeping vegetation, dry streambed, mixed 

hardwood-conifer (1 female); sweeping Sphagnum bog (1 male); sweeping understory shrubs, red 

maple stand (1 male); sweeping rhodora and Spiraea, mixed conifer-opening (1 female); sweeping 

meadowsweet, mixed conifer opening (1 male); sweeping grasses and forbs, school playground edge, 

disturbed area (3 males, 3 females); in nest on grasses, old fi eld (1 female); in silk retreat on black 

rush, saltmarsh (1 female); in silk retreats, red maple leaves (dead) on Juncus sp., brackish marsh 

(2 females); in nests, folded dead leaves impaled on black rush, brackish marsh (2 females); in retreat, 

impaled leaf on black rush, brackish marsh (1 female); in silk retreats, folded (dead) red maple leaves 

impaled on black rush stems, saltmarsh edge (4 females); in silk retreat, impaled leaf on rush, brackish 

marsh (1 female); on cobble beach with dipteran prey, seashore (1 female); on foliage in fl ower garden 

(1 female); in trash bin (1 male); n. d., old fi eld-brackish marsh edge (1 female); n. d., saltmarsh 

(1 male).

Regional Distribution: ME (Bryant 1908, as Evarcha hoyi (Peckham) Simon); CT (Kaston 1981, as 

Evarcha hoyi (Peckham)); NY (Crosby and Bishop 1928); QC (Bélanger and Hutchinson 1992).

Ghelna canadensis (Banks, 1897)

Taxonomy: Chickering (1944, as Metaphidippus canadensis (Banks)); Kaston (1973, 1981, as M. 

canadensis (Banks)); Maddison (1996); Paquin and Dupérré (2003).

Records: M12 (2 males, 1 female); M26 (1 female); M56 (9 males, 5 females). N = 18 adults; 11 

males, 7 females.

Methods: condensed litter-Berlese funnel (2 males); condensed litter-hand sorted (7 males, 

5 females); pitfall traps (1 male, 1 female); search (1 male, 1 female).

Months: June-July (1 female); July-August (1 male); August (1 male, 1 female); September (9 males, 

5 females).

Habitats: in nests, meadowsweet seedheads (1 male, 1 female); litter, freshwater pond edge (1 male); 

litter, saltmarsh edge (1 female); sifted litter (grasses, forbs, tidal debris), salt meadow (9 males, 5 

females).
146

Regional Distribution: ME (Mairs and Jennings, unpubl.); MA, NY, ON (Kaston 1973, 1981, as 

Metaphidippus canadensis (Banks)); QC (Bélanger and Hutchinson 1992).

Habronattus borealis (Banks, 1895)

Taxonomy: Chickering (1944); Kaston (1981); Griswold (1987); Paquin and Dupérré (2003).

Records: M04 (2 females); M08 (2 males, 5 females); M35 (1 female). N = 10 adults; 2 males, 

8 females.

Methods: search (1 male, 6 females); sweep net (1 male, 2 females).

Months: May (2 females); June (2 males, 5 females); July (1 female).

Habitats: sweeping marsh vegetation near brackish pond, seashore-brackish marsh (1 female); 

sweeping grasses and shrubs on shore, seashore-backshore (1 male, 1 female); among rocks at shore, 

seashore (1 male); in retreats under rocks, cobblestone beach, seashore (4 females); in silk retreat, red 

maple leaf on rush, brackish marsh (1 female); on cobblestone beach and ledge, seashore (1 female).

Regional Distribution: ME (Peckham and Peckham 1909, as Pellenes borealis B.); MA, NY, ON 

(Griswold 1987); CT (Kaston 1981); QC (Paquin et al. 2001).

Habronattus calcaratus maddisoni Griswold, 1987

Taxonomy: Griswold (1987); Paquin and Dupérré (2003).

Records: M04 (1 male); M58 (1 female). N = 2 adults; 1 male, 1 female.

Method: search (1 male, 1 female).

Month: May (1 male, 1 female).

Habitats: on rock ledge, seashore (1 male); on ground, marsh grass and matted straw, seashorebrackish marsh (1 female).

Regional Distribution: ME, MA, CT, NY, ON, QC (Griswold 1987).

Habronattus decorus (Blackwall, 1846)

Taxonomy: Chickering (1944); Kaston (1981); Griswold (1987); Paquin and Dupérré (2003).

Records: M01 (1 male); M06 (5 males); M35 (1 male). N = 7 adult males.

Methods: pitfall traps (5 males); search (1 male); sweep net (1 male).

Months: May (1 male); June (2 males); July (4 males).

Habitats: litter, saltmarsh (5 males); sweeping shrubs and grasses, brackish marsh (1 male); in garden 

(1 male).
147

Regional Distribution: ME (Procter 1933, as Pellenes splendens (Peckham)); NH, VT, MA, CT, NY, 

ON, NS (Griswold 1987); QC (Hutchinson 1999).

Habronattus viridipes (Hentz, 1846)

Taxonomy: Chickering (1944, as Habronattus peregrinus (Peckham), female only); Kaston (1981); 

Griswold (1987); Paquin and Dupérré (2003).

Records: M07 (2 males); M11 (1 female). N = 3 adults; 2 males, 1 female.

Habitats: litter, Kalmia-Vaccinium heath (2 males); ground, gravel pit in conifer-mixed hardwood 

opening (1 female).

Method: pitfall traps (2 males, 1 female).

Months: May (1 male); June (1 male); July (1 female).

Regional Distribution: ME (Peckham and Peckham 1909, as Pellenes peregrinus P.); VT, CT, NY, 

ON (Griswold 1987); QC (Bélanger and Hutchinson 1992).

**Marpissa grata (Gertsch, 1936)

Taxonomy: Gertsch (1936, as Hyctia grata Gertsch); Barnes (1958).

Record: M33 (1 female). N = 1 adult female.

Method: sweep net (1 female).

Month: June (1 female).

Habitat: sweeping grasses and sedges, freshwater marsh (1 female).

Regional Distribution: ME (this study).

Note: Previously known only from Minnesota and Michigan (Barnes 1958).

Neon nelli Peckham & Peckham, 1888

Taxonomy: Chickering (1944, as Neon nellii Peckham); Gertsch and Ivie (1955); Kaston (1981, as 

Neon nellii Peckham); Paquin and Dupérré (2003).

Records: M01 (1 female); M04 (1 male); M24 (1 female). N = 3 adults; 1 male, 2 females.

Method: pitfall traps (1 male, 2 females).

Months: June (1 male, 1 female); July-August (1 female).
148

Habitats: litter, seashore-backshore (1 female); litter, coastal red spruce stand (1 male); litter, red 

maple sapling stand (1 female).

Regional Distribution: ME (Bryant 1908); NH, MA, CT, NY, ON, NF, LB (Gertsch and Ivie 

1955); QC (Bélanger and Hutchinson 1992).

Pelegrina fl aviceps (Kaston 1973)

Taxonomy: Kaston (1973, 1981 as Metaphidippus fl aviceps Kaston); Maddison (1996); Paquin and 

Dupérré (2003).

Records: M01 (26 males, 13 females); M04 (4 males, 12 females); M06 (2 males, 3 females); M10 

(1 male, 3 females); M13 (1 male, 2 females); M15 (1 male, 2 females). N = 70 adults; 35 males, 35 

females.

Methods: beating cloth (34 males, 33 females); search (1 male); sweep net (2 females).

Months: April (1 male); May (20 males, 12 females); June (2 males, 2 females); August (9 males, 17 

females); September (1 female); October (3 males, 3 females).

Habitats: beating red spruce foliage, old fi eld edge (8 males, 4 females); beating red spruce foliage, 

old fi eld (3 males); beating red spruce foliage (10 males, 5 females); beating spruce foliage (branches) 

(4 males, 4 females); beating red spruce foliage, seashore-backshore (4 males, 11 females); beating 

spruce at shore (1 female); beating white spruce foliage, saltmarsh edge (2 males, 1 female); 

beating eastern larch foliage (saplings) (1 female); beating red spruce foliage (saplings), conifermixed hardwood (1 male, 2 females); beating red spruce foliage, mixed hardwood-conifer (1 male, 

2 females); beating red spruce foliage, saltmarsh edge (1 male, 2 females); sweeping vegetation, 

saltmarsh (1 female); sweeping grasses, saltmarsh (1 female); on red spruce (1 male).

Regional Distribution: ME, CT (Kaston 1973, 1981, as Metaphidippus fl aviceps Kaston); NH, VT, 

NY, ON, QC (Maddison 1996).

Note: Kaston’s (1973) record of Pelegrina fl aviceps in Maine refers to a collection by E. B. Bryant on 

4 August 1904 at Casco Bay, Cumberland County. He also noted that this species has been found at 

many other localities in Maine, but did not give specifi cs.

Pelegrina fl avipes (Peckham & Peckham, 1888)

Taxonomy: Chickering (1944, as Metaphidippus fl avipedes (Peckham)); Kaston (1973, 1981, as 

Metaphidippus fl avipedes (Peckham)); Maddison (1996); Paquin et al. (2001) and Paquin and 

Dupérré (2003).

Records: M01 (4 males, 10 females); M04 (5 males, 3 females); M05 (1 male); M06 (3 females); 

M15 (2 males, 2 females); M17 (1 female); M32 (2 females); M38 (1 female); M48 (1 male, 

1 female); M49 (3 males). N = 39 adults; 16 males, 23 females.

Methods: beating cloth (15 males, 20 females); sweep net (1 male, 3 females).
149

Months: May (1 male, 4 females); June (2 males, 8 females); July (3 males, 2 females); August 

(7 males, 6 females); September (3 males, 2 females); October (1 female).

Habitats: beating red spruce foliage, old fi eld (2 males, 1 female); beating red spruce foliage, old 

fi eld edge (1 female); beating red spruce foliage, upper shore, seashore-backshore (2 females); beating 

red spruce foliage (1 male, 3 females); beating spruces (1 female); beating white pine foliage (1 

male, 1 female); beating spruce at shore (5 males, 3 females); beating white spruce foliage, saltmarsh 

edge (3 females); beating red spruce foliage, saltmarsh edge (2 males, 2 females); beating red spruce 

foliage, mixed hardwood-conifer (2 females); beating red spruce foliage, seashore-backshores (4 male, 

1 female); sweeping Vaccinium and small shrubs, old fi eld (1 female); sweeping wet meadow, old 

fi eld-brackish marsh (1 male); sweeping grasses, saltmarsh (1 female); sweeping Sphagnum bog 

(1 female).

Regional Distribution: ME (Bryant 1908, as Dendryphantes fl avipedes Peckham); CT (Kaston 1981, 

as Metaphidippus fl avipedes (Peckham)); NY (Crosby and Bishop 1928, as Metaphidippus fl avipedes 

(Peck.); NB (Loughton et al. 1963, as Metaphidippus fl avipedes (Peckham)); ON, QC, NS, PE, NF 

(Maddison 1996).

Pelegrina insignis (Banks, 1892)

Taxonomy: Kaston (1981, as Metaphidippus insignis (Banks)); Maddison (1996); Paquin and 

Dupérré (2003).

Records: M01 (3 males, 12 females); M05 (3 males, 11 females); M06 (2 males, 8 females) M07 (1 

female); M08 (1 female); M12 (2 males, 8 females); M30 (1 female); M33 (1 male, 3 females); M35 

(4 females); M41 (1 female); M47 (3 females); M51 (1 female); M55 (4 males, 3 females). N = 72 

adults; 15 males, 57 females.

Methods: search (4 females); sweep net (15 males, 53 females).

Months: May (4 males, 12 females); June (11 males, 31 females); July (11 females); August (3 

females).

Habitats: sweeping vegetation, old fi eld (3 males, 10 females); sweeping grasses and forbs in 

open fi eld, old fi eld (1 male, 9 females); sweeping wet meadow, old fi eld-brackish marsh (2 males, 

2 females); sweeping vegetation (Juncus-Spartina), saltmarsh (1 male, 8 females); sweeping grasses 

and low shrubs, saltmarsh edge (1 male); sweeping grasses and shrubs on shore, seashore-backshore 

(1 female); sweeping marsh grasses and shrubs (Spiraea-Kalmia), freshwater marsh (2 males, 2 

females); sweeping grasses, freshwater marsh (2 females); sweeping shrubs, freshwater marsh edge 

(3 females); sweeping vegetation (1 female); sweeping grasses, forbs, ferns, and rushes, roadside 

alder-aspen edge (1 female); sweeping marsh grasses and sedges, freshwater marsh (1 male, 3 

females); sweeping vegetation, brackish marsh (4 females); sweeping vegetation, freshwater streamside 

(3 females); sweeping rhodora and Spiraea, mixed conifer opening (1 female); sweeping grasses and 

forbs, school playground edge, disturbed area (4 males, 3 females); in web, head of grass stem, with 

remains of Dictyna coloradensis and various Diptera, old fi eld (1 female); in dry, folded red maple 

leaf, saltmarsh edge (1 female); on heath plants, Kalmia-Vaccinium heath (1 female); n. d., old fi eld 

(1 female).
150

Regional Distribution: ME (Richman and Cutler 1978, as Metaphidippus insignis (Banks); VT, MA, 

CT, NY, NB, ON (Maddison 1996); QC (Bélanger and Hutchinson 1992).

Pelegrina proterva (Walckenaer, 1837)

Taxonomy: Kaston (1973, 1981, as Metaphidippus proterva (Walckenaer); Maddison (1996); Paquin 

and Dupérré (2003).

Records: M01 (8 males, 11 females); M08 (2 females); M12 (1 male, 2 females); M14 (1 female); 

M15 (1 male); M24 (1 female); M26 (4 males, 2 females); M35 (1 female); M37 (1 male, 2 females); 

M47 (1 female). N = 38 adults; 15 males, 23 females.

Methods: beating cloth (1 male); pitfall trap (1 female); search (1 male, 2 females); sifted litter-hand 

sorted (1 male); sweep net (12 males, 20 females).

Months: May (1 male, 1 female); June (9 males, 15 females); July (4 females); August (3 females); 

September (4 males); October (1 male).

Habitats: beating foliage in alder thicket (1 male); sifting forest-fl oor litter under maple-birch 

overstory, mixed hardwood-conifer (1 male); litter, saltmarsh edge (1 female); sweeping vegetation, 

old fi eld (5 males, 3 females); sweeping vegetation, old fi eld edge (4 females); sweeping asters, 

goldenrod, and small alders, old fi eld (1 male); sweeping meadowsweet stand (1 female); sweeping 

grasses on shore, seashore-backshore (1 female); sweeping grasses, forbs, and seaside-pea, seashoreupper beach (1 female); sweeping grasses and shrubs on shore, seashore-upper beach (1 female); 

sweeping marsh grasses and sedges, freshwater marsh (1 female); sweeping shrubs, freshwater marsh 

(1 female); sweeping grasses and forbs, mixed hardwood-conifer (1 female); sweeping grasses, 

saltmarsh edge (1 male); sweeping understory vegetation, red maple sapling stand (1 female); 

sweeping and shaking chokecherry foliage and fl owers, saltmarsh edge (4 males, 1 female); sweeping 

shrubs, brackish marsh (1 female); sweeping grasses and Spiraea, mown fi eld edge (1 male, 2 females); 

sweeping marsh vegetation, freshwater streamside (1 female); among alder leaves (1 female); in silk 

retreat with egg sac, folded Spiraea leaf (1 female); on rhodora in blossom, freshwater marsh (1 male).

Regional Distribution: ME (Procter 1938, as Dendryphantes capitatus (Hentz)); NH, VT, MA, CT, 

RI, NY, ON, QC, NS (Maddison 1996).

Phidippus clarus Keyserling, 1885

Taxonomy: Chickering (1944); Kaston (1981); Paquin and Dupérré (2003); Edwards (2004).

Records: M01 (11 males, 18 females); M05 (2 males, 3 females); M06 (5 females); M07 (1 male, 

3 females); M22 (1 male); M28 (2 males, 1 female); M35 (9 males, 6 females); M47 (1 male); M55 

(4 males, 4 females). N = 71 adults; 31 males, 40 females.

Methods: pitfall trap (1 male); search (3 males, 8 females); sweep net (27 males, 30 females); n. d. 

(2 females).

Months: June (4 males, 4 females); June-July (1 male); July (21 males, 15 females); August (5 males, 

15 females); September (6 females).
151

Habitats: litter, bigtooth aspen stand (1 male); sweeping vegetation, old fi elds (9 males, 12 females); 

sweeping vegetation, old fi eld edge (1 female); sweeping Vaccinium and Myrica near woods, old 

fi eld (1 male, 1 female); sweeping vegetation, saltmarsh (2 females); sweeping vegetation, KalmiaVaccinium heath (1 male, 3 females); sweeping white and yellow clover, roadside disturbed area 

(2 males, 1 female); sweeping vegetation, brackish marsh (9 males, 6 females); sweeping marsh 

vegetation, freshwater streamside, riparian (1 male); sweeping grasses and forbs, school playground 

edge, disturbed area (4 males, 4 females); in folded blackberry leaf (1 female); in silken nest, low 

shrub leaves (1 female); in leaf retreat, chokecherry seedling, old fi eld (1 female); in nest with egg sac, 

curled chokecherry leaves, old fi eld (1 female); in nest between two hawthorn leaves (1 female); in 

nest with spiderlings, hawthorn leaves (1 female); in retreats, goldenrod basal leaves (2 females); on 

wild cherry leaf (1 male); on twig, small pin cherry (1 male); running over ledge at shore, seashore 

(1 male); n. d., old fi eld (1 female); n. d., saltmarsh (1 female).

Regional Distribution: ME (Bryant 1908, as Phidippus multiformis Emerton); QC (Bélanger and 

Hutchinson 1992); NH, VT, MA, CT, RI, NY, ON (Edwards 2004).

Phidippus purpuratus Keyserling, 1885

Taxonomy: Chickering (1944); Kaston (1981); Paquin and Dupérré (2003); Edwards (2004).

Records: M01 (1 male); M04 (1 male, 1 female); M07 (1 female); M11 (1 male); M15 (1 female); 

M45 (1 male). N = 7 adults; 4 males, 3 females.

Method: search (4 males, 3 females).

Months: May (1 male, 1 female); June (2 males); July (1 male); August (2 females).

Habitats: among rocks, roadside embankment (1 male); in silk retreat under rock, coastal sprucemixed hardwood (1 female); in egg retreat under rock, Kalmia-Vaccinium heath (1 female); on 

Cladina-covered ledges, mixed conifer-hardwood (1 male); on cobblestone beach and ledge (1 male); 

on rock, gravel pit, mixed conifer-hardwood opening (1 male); on small rock, saltmarsh (1 female).

Regional Distribution: ME (Keyserling 1885); NH, VT, MA, CT, NY, ON, QC (Edwards 2004).

Phidippus whitmani Peckham & Peckham, 1909

Taxonomy: Chickering (1944); Kaston (1981); Paquin and Dupérré (2003); Edwards (2004).

Records: M01 (2 males); M27 (1 male). N = 3 adult males.

Method: search (3 males).

Month: June (3 males).

Habitats: in fl ower garden (1 male); on ground, red maple-white (paper) birch stand (1 male); inside 

window, house (1 male).

Regional Distribution: ME (Procter 1946); NH, MA, CT, RI, NY, ON, QC, NS (Edwards 2004).
152

Salticus scenicus (Clerck, 1757)

Taxonomy: Chickering (1944); Kaston (1981, as Salticus scenicus (Linnaeus)); Paquin and Dupérré 

(2003).

Records: M01 (8 males, 1 female); M21 (1 female). N = 10 adults; 8 males, 2 females.

Method: search (8 males, 2 females).

Months: May (2 males); June (6 males, 2 females).

Habitats: in wood bin (1 male); on ground, old fi eld edge (1 male); on ground, vegetable garden 

(1 male); on exterior walls, house (3 males, 1 female); on fl oor, house (1 male); on exterior of screen 

door, house (1 male); on exterior white vinyl siding, building (1 female).

Regional Distribution: ME (Bryant 1908); CT (Kaston 1981, as Salticus scenicus (Linnaeus)); NY 

(Crosby and Bishop 1928); QC (Bélanger and Hutchinson 1992).

**Sibianor aemulus (Gertsch, 1934)

Taxonomy: Gertsch (1934, as Sassacus aemulus n. sp.); Maddison (1978, as Bianor aemulus (Gertsch, 

1934)); Logunov (2001).

Records: M01 (2 males); M14 (1 male). N = 3 adult males.

Methods: pitfall traps (2 males); search (1 male).

Months: June (1 male); July (2 males).

Habitats: litter, deciduous-coniferous woodland (1 male); litter, Betula-Acer stand (1 male); on 

ground among grasses and forbs, mixed hardwood-conifer overstory (1 male).

Regional Distribution: ME (this study); ON (Richman and Cutler 1978, as Bianor aemulus 

(Gertsch); QC (Bélanger and Hutchinson 1992).

Note: Richman et al. (2005) also noted this species in Maine, but their record originated from this 

study.

Sitticus fl oricola palustris (Peckham & Peckham, 1883)

Taxonomy: Chickering (1944); Prószyński (1980); Kaston (1981, as Sitticus palustris (Peckham)); 

Paquin and Dupérré (2003).

Records: M01 (2 males); M05 (1 female); M06 (5 males, 2 females); M12 (1 male, 10 females); M15 

(1 female); M17 (3 males); M23 (1 female); M33 (1 male, 22 females); M34 (1 male, 1 female); M35 

(3 males, 27 females); M38 (1 male); M41 (1 male, 6 females); M47 (2 males, 3 females); M55 (2 

males, 2 females); M57 (1 female). N = 99 adults; 22 males, 77 females.
153

Methods: litter condenser-Berlese funnel (1 male); search (3 males, 65 females); sweep net (18 males, 

10 females); n. d. (2 females).

Months: June (5 males, 44 females); July (5 males, 25 females); August (12 males, 8 females).

Habitats: sifted grass litter near freshwater stream, riparian (1 male); sweeping foliage, freshwater 

pond edge (1 male); sweeping vegetation, old fi eld (1 female); sweeping vegetation (Juncus-Spartina), 

saltmarsh (1 male); sweeping grasses, saltmarsh (1 female); sweeping grasses and shrubs, saltmarsh 

(1 female); sweeping vegetation, saltmarshes (6 males); sweeping grasses and shrubs (Spiraea-Kalmia), 

freshwater marsh (1 male, 1 female); sweeping grasses, freshwater marshes (1 male, 1 female); 

sweeping grasses and sedges, freshwater marsh (1 female); sweeping grasses, sedges, and forbs, dry 

stream bed, mixed hardwood-conifer (1 male, 1 female); sweeping grasses, brackish marsh (2 females); 

sweeping vegetation, brackish marsh (3 males); sweeping vegetation, Sphagnum-cranberry-rush bog 

(1 male); sweeping marsh vegetation, freshwater streamside, riparian (2 males); sweeping grasses 

along freshwater stream, riparian (1 female); sweeping mixed vegetation (grasses, forbs) near school 

playground, disturbed area (1 male); in silk retreat, folded Spiraea leaf, freshwater pond edge (1 

male); in silk retreat on black rush (1 male); in silken retreats on meadowsweet, freshwater marsh 

(4 females); in retreats on rhodora (2 females); in retreat, fruiting head of rush, freshwater marsh 

(1 female); in silk retreats, dry infl orescences of meadowsweet, freshwater marsh (21 females); in 

adjacent nests, rolled leaves stuck on black rush, brackish marsh (3 females); in retreats, leaves and 

seedhead of marsh grass, brackish marsh (3 females); in retreats, dry composite seedhead, brackish 

marsh (4 females); in silk retreats, red maple leaf on Juncus sp., brackish marsh (2 females); in retreats, 

dry forb stem, brackish marsh (4 females); in retreats, impaled maple leaves on black rush stems, 

brackish marsh (5 females); in nests, folded dead leaves on black rush, brackish marsh (3 females); in 

nest with young spiderlings on Juncus (1 female); in silk retreats, dead red maple leaves impaled on 

black rush stems, saltmarsh edge (1 male, 6 females); in silk retreats on Scirpus seedhead, freshwater 

marsh, riparian (3 females); in retreat, dry infl orescence of meadowsweet, freshwater streamside 

(1 female); in silk retreat, impaled tree leaf on black rush (1 female); on ground under paper, bog 

edge, island (1 female); n. d., saltmarsh (1 female); n. d., freshwater marsh (1 female).

Regional Distribution: ME (Emerton 1891, as Attus palustris Peckham); CT (Kaston 1981, as 

Sitticus palustris (Peckham)); NY (Crosby and Bishop 1928); QC (Bélanger and Hutchinson 1992, as 

Sitticus palustris (Peckham & Peckham)).

Note: Most North American investigators of Salticidae consider Prószyński’s (1980) designation of S. 

palustris as a subspecies of S. fl oricola (C. L. Koch, 1837), a European species, to be invalid (B. Cutler, 

pers. comm.). However, the basis for such emendation remains to be made; see Platnick (2007).

*Sitticus pubescens (Fabricius, 1775)

Taxonomy: Roberts (1985, 1987); Kaston (1981).

Record: M21 (1 male). N = 1 adult male.

Method: search (1 male).
154

Month: August (1 male).

Habitat: on exterior white vinyl siding, building (1 male).

Regional Distribution: ME (this study); MA, CT (Kaston 1981).

Note: Sitticus pubescens is an invasive species from Europe, which was fi rst discovered in North 

America at Allston, MA (Bryant 1941). It has since been found in Connecticut (Kaston 1981) and in 

New Jersey (B. Cutler, pers. comm.). 

Sitticus striatus Emerton, 1911

Taxonomy: Prószyński (1980); Kaston (1981); Paquin and Dupérré (2003).

Records: M06 (1 male, 1 female); M56 (2 females). N = 4 adults; 1 male, 3 females.

Methods: litter condenser-hand sorted (2 females); pitfall trap (1 female); sweep net (1 male).

Months: July (1 female); August (1 male); September (2 females).

Habitats: litter, saltmarsh (1 female); sifted litter (grasses, forbs, tidal debris), salt meadow 

(2 females); sweeping vegetation, saltmarsh (1 male).

Regional Distribution: ME (Emerton 1911, as Sittacus [sic.] striatus Emerton); CT (Kaston 1981); 

MA, NF (Richman and Cutler 1978); QC (Bélanger and Hutchinson 1992); LB (Prószyński 1980).

*Synageles noxiosus (Hentz, 1850)

Taxonomy: Kaston (1981); Cutler (1987); Paquin and Dupérré (2003).

Record: M12 (1 male). N = 1 adult male.

Method: sweep net (1 male).

Month: June (1 male).

Habitat: sweeping grasses and other vegetation, freshwater marsh (1 male).

Regional Distribution: ME (this study); MA, CT, NY, ON (Cutler 1987); QC (Hutchinson 1999).

Talavera minuta (Banks, 1895)

Taxonomy: Kaston (1981); Paquin and Dupérré (2003).

Records: M06 (1 male); M07 (1 female). N = 2 adults; 1 male, 1 female.

Methods: pitfall trap (1 male); unsifted litter-Berlese funnel (1 female).
155

Months: July (1 male); August (1 female).

Habitats: litter, saltmarsh (1 male); litter, Kalmia-Vaccinium heath (1 female).

Regional Distribution: ME (Collins et al. 1996); MA, CT, NY (Richman and Cutler 1978); QC 

(Bélanger and Hutchinson 1992).

Tutelina similis (Banks, 1895)

Taxonomy: Kaston (1981); Paquin and Dupérré (2003).

Records: M51 (2 females); M55 (2 females). N = 4 adult females.

Method: sweep net (4 females).

Month: June (4 females).

Habitats: sweeping meadowsweet, mixed conifer opening (1 female); sweeping rhodora and 

meadowsweet, mixed conifer opening (1 female); sweeping grasses and forbs, school playground edge, 

disturbed area (2 females).

Regional Distribution: ME (Procter 1938, 1946, as Icius similis Banks); CT (1981); NY (Crosby 

and Bishop 1928, as Icius similis Banks); QC (Bélanger and Hutchinson 1992).
156

FAUNAL ATTRIBUTES AND HABITAT ASSOCIATIONS

Spider Taxa

Adult spiders of 19 families, 145 genera, and 302 species (4 unknown) were collected from diverse 

habitats in Milbridge, Washington County, ME, during 1991-2005. By comparison, Procter (1946) 

listed spiders of 15 families, 94 genera, and 179 species from various habitats of Mount Desert 

Island, Hancock County, ME. With recent taxonomic revisions, Procter’s faunal list now represents 

22 families, 109 genera, and 179 species (2 unknown). Studies of the ground-inhabiting fauna 

of blueberry fi elds in Washington County (Milbridge excluded) yielded spiders of 17 families, 53 

genera, and 87 species (Collins et al. 1996). Maloney (2002) also investigated the terrestrial spider 

fauna of blueberry fi elds in Washington County and reported 17 families, 81 genera, and 133 species. 

In Maine’s northern spruce-fi r forests, a pitfall-trap study yielded spiders of 15 families, 76 genera, 

and at least 125 species associated with forest-fl oor litter (Jennings et al. 1988). Thus, comparatively, 

the Milbridge study represents the most comprehensive inventory of spiders for any locale within 

Maine, and includes taxa associated with diverse communities, habitats, and strata.

Species richness per family at Milbridge ranged from 1 to 89; overall mean = 15.9 ± 4.6, N = 19. Not 

surprisingly, the Linyphiidae were represented by the greatest number of genera and species, with 19 

genera and 35 species of the subfamily Linyphiinae, and 28 genera and 54 species (2 unknown) of 

the subfamily Erigoninae. These sheet-web spiders comprise diverse assemblages of genera and species 

in New England (Hilburn and Jennings 1988, Jennings et al. 1988, Kaston 1981) and in eastern 

Canada (Bélanger and Hutchinson 1992, Paquin et al. 2001). Conversely, three spider families at 

Milbridge were represented by only a single species each: the Pholcidae, Titanoecidae, and Pisauridae. 

All three families have a limited number of species per family in the northern region, i.e., two species 

of Pholcidae, one species of Titanoecidae, and six species of Pisauridae (Dondale and Redner 1990, 

Kaston 1981, Paquin and Dupérré 2003).

Species richness per genus at Milbridge ranged from 1 to 13; overall mean = 2.1 ± 0.2, N = 145. Fully 

88 genera (60.1 percent of total genera) of spiders collected at Milbridge were represented by a single 

species; most were genera of Linyphiinae and Erigoninae. However, some of the collected genera 

have only one known species in North America (e.g., Neottiura, Estrandia, Macrargus, Metopobactrus, 

and Trabeops); others have limited numbers of species in New England (e.g., Allomengea, Tapinopa, 

Misumena, Misumenoides, Sibianor, and Ghelna). The fi ve richest genera at Milbridge were Clubiona, 

N = 13 species; Xysticus, N = 12 species; Pardosa, N = 10 species; Walckenaeria, N = 9 species; and 

Theridion, N = 8 species.

Spider Numbers

Fully 6,979 adult spiders were collected during this inventory. In addition, 3,980 juvenile spiders 

were also taken but were discarded because of their limited contribution to species richness. Few 

juvenile spiders can be determined to species with certainty. Most of the juvenile spiders were 

cursorial species captured by pitfall traps. As expected, adult spider abundances were unequally 

distributed among families, genera, and species. Individual specimens per family ranged from 1 

(Titanoecidae, Pisauridae) to 1,691 (Lycosidae) (Fig. 2); overall mean = 367.3 ± 97.9, N = 19. 

Individual specimens per genus ranged from 1 (N = 22 genera) to 1,159; overall mean = 48.1 ± 9.2, 

N = 145. The fi ve most frequently collected genera were Pardosa, N = 1,159; Neoantistea, N = 314; 

Clubiona, N = 248; Xysticus, N = 245; and Trochosa, N = 222. Individual specimens per species ranged 
157

from 1 to 470; overall mean = 23.1 ± 2.7, N = 302. Fully 53 species were represented by a single 

specimen.

The 10 most frequently collected species of spiders at Milbridge were Pardosa moesta Banks, 1892, 

N = 470; P. modica (Blackwall, 1846), N = 326; Neoantistea magna (Keyserling, 1887), N = 283; 

Eris militaris (Hentz, 1845), N = 207; Zelotes fratris Chamberlin, 1920, N = 172; P. xerampelina 

(Keyserling, 1877), N = 170; Enoplognatha ovata (Clerck, 1757), N = 158; Neoscona arabesca 

(Walckenaer, 1842), N = 156; Trochosa terricola Thorell, 1856, N = 156; and Grammonota angusta 

Dondale, 1959, N = 137. Collectively, these 10 species represented about one-third (32.0 percent) of 

the total adult spiders (N = 6,979) taken during this inventory.

Spider Foraging Guilds

Spider taxa and abundance of individuals also varied by two basic foraging strategies or guilds (Root 

1973, Uetz et al.1999), i.e., web spinners and hunters. Web-spinning spiders spin webs to capture 

their prey, whereas hunting spiders rely on visual cues and use stealth, ambush, and chase to capture 

prey. More families, genera, and species of web spinners were collected than hunters. Of the 19 

families of spiders collected at Milbridge, 10 belonged to the web spinner guild and 9 to the hunter 

guild. Genera of web spinners (N = 98) per guild family ranged from 1 (Pholcus, Titanoeca) to 49 

(Linyphiinae, Erigoninae); overall mean = 9.8 ± 4.6, N = 10. Genera of hunters (N = 47) per guild 

family ranged from 1 (Dolomedes, Agroeca, Clubiona) to 15 (Salticidae); overall mean = 5.2 ± 1.6, N = 

9. Species of web spinners (N = 179) per guild family ranged from 1 (Pholcus manueli Gertsch, 1937; 

0

200

400

600

800

1,000

1,200

1,400

1,600

1,800

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19

Spider Family

Number of Spiders

Lycosidae

Linyphiidae

Salticidae

Theridiidae

Araneidae

Hahniidae

Thomisidae

Gnaphosidae

Clubionidae

Philodromidae

Agelenidae

Corinnidae

Tetragnathidae

Amaurobiidae

Dictynidae

Liocranidae

Pholcidae

Titanoecidae

Pisauridae

Hunters

Web Spinners

Figure 2.—Rank-order abundances of spider families collected in Milbridge, Washington County, ME, during the 

period 1991-2005.
158

Titanoeca americana Emerton, 1888) to 89 (Linyphiinae, Erigoninae); overall mean = 17.9 ± 8.4, N = 

10. Species of hunters (N = 123) per guild family ranged from 1 (Dolomedes striatus Giebel, 1869) to 

25 (Lycosidae, Salticidae); overall mean = 13.7 ± 2.9, N = 9. Both the Lycosidae and Salticidae were 

represented by 25 species each.

The preponderance of web spinner taxa over hunter taxa in the Milbridge inventory was not 

unexpected. Kaston (1981) noted there were more families, genera, and species of web spinners than 

hunters in all of New England. Excluding those families not found at Milbridge, Kaston’s totals 

were for web spinners, 146 genera and 376 species; for hunters, 73 genera and 278 species. The 

proportional representation of these guilds at Milbridge was remarkably similar to those estimated by 

Kaston for New England. Web spinners made up 67.6 percent of the total genera (N = 145) and 59.3 

percent of the total species (N = 302) observed at Milbridge; the corresponding estimated values for 

web spinners in New England were 66.7 percent genera and 57.5 percent species. Conversely, hunters 

made up 32.4 percent of the total genera, and 40.7 percent of total species observed at Milbridge; 

the corresponding estimated values for hunters in New England were 33.3 percent genera and 

42.5 percent species. G-tests indicated that none of the observed vs. estimated proportions differed 

signifi cantly by guild; for genera, G < 0.005, P ≤ 0.05; for species, G = 0.40, P ≤ 0.05.

During the 15-year study period, more hunters (N = 3,806) were collected than web spinners (N = 

3,173). Among web spinners, the number of individuals per family ranged from 1 (Titanoecidae) 

to 1,097 (Linyphiidae); guild mean = 317.3 ± 106.4, N = 10. Among hunters, the number of 

individuals per family ranged from 1 (Pisauridae) to 1,691 (Lycosidae); guild mean = 422.9 ± 175.2, 

N = 9. Collectively, both guilds were represented by 6,979 spiders; overall guild mean = 367.3 ± 97.9, 

N = 19 families.

Spider guild representation also differed by genera and by species. Among web spinners, the number 

of individuals per genus ranged from 1 (14 genera) to 314 (Neoantistea); guild mean = 32.4 ± 5.3, 

N = 98; among hunters, the number of individuals per genus ranged from 1 (4 genera) to 1,159 

(Pardosa); guild mean 81.0 ± 25.7. For both foraging guilds, overall mean = 48.1 ± 9.2 individuals per 

genus, where N = 145.

Collected individuals per web spinner species ranged from 1 (38 species) to 283 (Neoantistea magna 

(Keyserling, 1887)); overall guild mean = 17.7 ± 2.6, N = 179. Collected individuals per hunter 

species ranged from 1 (15 species) to 470 (Pardosa moesta Banks, 1892). For both guilds, overall mean 

= 23.1 ± 2.7 individuals per species, where N = 302.

Spider Sex Ratios

More female spiders (N = 3,870) were collected than male spiders (N = 3,109); overall female: male 

ratio = 1.24: 1.00. Not surprisingly, spider sex ratios varied widely among collected species and by 

foraging guild, and were infl uenced by sampling method, habitat structure, spider life histories, and 

seasonal differences.

For both foraging guilds, the proportional distribution of the sexes among species varied from 0.0 

percent to 100.0 percent, with web spinners contributing more to female biasness than hunters. 

Among the 179 species of web spinners, 109 (60.9 percent) were female biased, 54 (30.2 percent) 

were male biased, and 16 (8.9 percent) reached unity (50.0 percent female, 50 percent male). By 
159

contrast, among the 123 species of hunters, 61 (49.6 percent) were female biased, 52 (42.3 percent) 

were male biased, and 10 (8.1 percent) reached unity (50.0 percent female, 50.0 percent male).

Among web spinner species (N = 179), collected males per species ranged from 0 to 247 (Neoantistea 

magna (Keyserling, 1887)), mean = 7.6 ± 1.6; collected females per species ranged from 0 to 128 

(Enoplognatha ovata (Clerck, 1757)), mean = 10.2 ± 1.5. Among hunter species (N = 123), collected 

males per species ranged from 0 to 219 (Pardosa moesta Banks, 1892), mean = 14.3 ± 2.5; collected 

females per species ranged from 0 to 251 (Pardosa moesta), mean = 16.7 ± 3.2. For both guilds 

combined (N = 302 species), the overall means per species were 12.8 ± 1.6 females, and 10.3 ± 1.4 

males.

Three distributional patterns were evident among these spider guild-sex comparisons: 1) within 

guilds, mean abundances were consistently higher for females than males; 2) between guilds, mean 

abundances of both sexes were consistently higher for hunters than for web spinners; and 3) for both 

guilds combined, mean abundances were consistently higher for females than males.

Of the 179 web spinner species, 44 (24.6 percent) were represented by females only; 25 (14.0 

percent) were represented by males only; both sexes made up the remaining 110 (61.4 percent) 

species of web spinners.

Most (58.0 percent, N = 69) of the web spinner species represented by only one sex were linyphiids 

(N = 40 species), especially small erigonids of the subfamily Erigoninae (N = 30). However, species 

of other families were also included in the single sex category; 12 species of theridiids, 3 species of 

tetragnathids, 8 species of araneids, 5 species of dictynids, and 1 species of Titanoecidae. For female 

only species, the number of collected individuals ranged from 1 to 114; for male only species, from 1 

to 4.

Collections of two species of web spinners, Hypselistes fl orens (O. Pickard-Cambridge, 1875) and 

Hypsosinga pygmaea (Sundevall, 1831), yielded signifi cant numbers of females but no males: Hypselistes 

fl orens, N = 114 females; Hypsosinga pygmaea, N = 54 females. Both species mature in early spring; 

males of Hypselistes fl orens have been taken through June in Connecticut (Kaston 1981); males of 

Hypsosinga pygmaea in May and June in Minnesota and elsewhere (Levi 1971). Although numerous 

sweep-net samples were taken during May and June at Milbridge, none yielded males of either species.

Of the 123 hunter species, 13 (10.6 percent) were represented by females only; 19 (15.4 percent) were 

represented by males only; both sexes made up the remaining 91 (74.0 percent) species of hunters.

Most of the hunter species represented by only one sex (N = 32) were distributed unequally among 

eight families: six species of Lycosidae, one species of Pisauridae, fi ve species of Clubionidae, two 

species of Corinnidae, one species of Gnaphosidae, four species of Philodromidae, fi ve species of 

Thomisidae, and eight species of Salticidae.

Sampling methodology and habitat structure contributed to the dominance of female spiders 

collected at Milbridge. For example, female-occupied webs of araneids and linyphiids were visibly 

prominent in the herb-shrub-small tree strata of old fi elds, marshes, and forests, especially during 

and immediately after coastal fogs. Searches and sweep-netting of these strata and habitats yielded 
160

numerous females of Araneus marmoreus Clerck, 1757 and Frontinella communis (Hentz 1850). 

Beating coniferous-tree foliage yielded chiefl y females of Theridion murarium Emerton, 1882; T. 

pictum Walckenaer, 1802; Pityohyphantes costatus (Hentz, 1850); and P. phrygianus (C. L. Koch, 

1836). Pitfall trapping in the litter layer of saltmarshes yielded males of Pardosa modica (Blackwall, 

1846) early on; however, as the season progressed, females were more abundant among trap 

catches. Sweeping herbaceous vegetation of old fi elds and roadsides yielded numerous females of 

Enoplognatha ovata (Clerck, 1757), but few males. Females but no males of Hypselistes fl orens (O. 

Pickard-Cambridge, 1875) were also swept from old fi eld and roadside vegetation. Females but no 

males of Hypsosinga pygmaea (Sundevall, 1831) were swept chiefl y from herbaceous vegetation of salt, 

brackish, and freshwater marshes. Several other examples of female-biased collections were evident 

and attributable to sampling methodology and habitat.

Spider life histories no doubt contributed to the apparent disparity between the collected sexes at 

Milbridge. Male spiders generally have fewer molts and reach adulthood before the females (Foelix 

1996, Gertsch 1979, Schaefer 1987). Male spiders also have shorter lifespans than the females; hence, 

sex ratios of fi eld-collected spiders are usually characterized by fewer males (Schaefer 1987), especially 

as the season progresses. With few exceptions, this was the case at Milbridge. For example, adult 

males of some lycosid species were captured early in the season before adult females appeared in the 

same traps. Two notable exceptions to this protandrous pattern of activity were evidenced by pitfalltrap captures of the hahniid Antistea brunnea (Emerton 1909) and the corrinid Castianeira cingulata

(C. L. Koch 1841). Females of A. brunnea were taken in June and July, whereas males were captured 

in August and September, chiefl y the latter. Females of C. cingulata were taken monthly from May to 

August; males were captured only in August.

Another factor that no doubt contributed to the greater abundance of female spiders among the 

Milbridge collection was our tendency to collect females during their reproductive period. Because of 

their larger size and consequent greater visibility to observers, female lycosids with egg sacs were more 

apt to be collected than the smaller males. Females of Pardosa xerampelina (Keyserling 1877) with egg 

sacs were collected more frequently than the males of this species. Multiple collections of Clubiona 

riparia L. Koch, 1866 and Sitticus fl oricola palustris (Peckham & Peckham 1883) females also were 

made, once their nesting microhabitats had been discovered. In fresh and saltwater marshes, females 

of C. riparia construct their nests by folding Spartina leaves into an oblong, three-sided retreat; see 

Comstock (1948, fi g. 643). Such retreats are easily detected once a search image has been formed. At 

the edges of freshwater marshes and elsewhere, females of Sitticus fl oricola palustris spin retreats on the 

dried infl orescences of meadowsweet. Retreats of this salticid spider were often found in aggregations 

of four to fi ve females per infl orescence, with one female per retreat. These examples of observerbiased collections contributed to the disparity between the sexes.

Collection Method Effects

Pitfall traps yielded more species of spiders (N = 179) and more spider individuals (N = 3,143) than 

any other sampling method used at Milbridge (Figs. 3 and 4). This result was not surprising because 

we used pitfall traps to sample the ground-inhabiting fauna at several different sites and habitats. 

Once deployed, the traps were operated over extended periods of time (months), thus yielding both 

diurnally active and nocturnally active species. Because some spiders exhibit diel activity patterns 

(Williams 1962), several investigators have stressed the importance of sampling both during the day 

and at night (Green 1999, Sørensen et al. 2002, and others).
161

0

20

40

60

80

100

120

pitfall trap search sweep net beating sheet litter extraction other

Collection Method

Number of Species

Hunters

Web Spinners

Figure 3.—Number of species of spiders by collection method. Other includes aquatic pitfall traps, brushing treebole bark, shaking foliage over net, and no data.

Figure 4.—Number of spiders by collection method. Other includes aquatic pitfall traps, brushing tree-bole bark, 

shaking foliage over net, and no data.

0

200

400

600

800

1,000

1,200

1,400

1,600

1,800

2,000

2,200

pitfall trap sweep net search beating sheet litter extraction other

Collection Method

Number of Spiders

Hunters

Web Spinners
162

Although pitfall-trapping yielded the most species and most individuals, searches, sweep-netting, 

and beating-cloth collections were also productive at Milbridge; searches yielded 1,545 spiders of 

155 species; sweep-netting yielded 1,470 spiders of 132 species; beating foliage, chiefl y of deciduous 

and coniferous trees, yielded 586 spiders of 56 species. Each of the remaining sampling methods 

(brushing tree-bole bark, sifted and unsifted litter-Berlese funnel, sifted litter-hand sorted, shaking 

foliage-sweep net, and aquatic pitfalls) used at Milbridge yielded < 75 individuals and < 35 species. 

Nonetheless, the importance of these low-yielding methods was evidenced by the number of methodunique species (see below).

Except for aquatic pitfall traps which yielded only one species of hunting spider, species of web 

spinners were consistently captured more frequently than hunters regardless of method (Fig. 3). This 

dominant web spinner trend held true even for the lesser yielding methods, such as brushing bark, 

sifting litter, and shaking foliage. However, the abundance of individual spiders was more variable 

among methods (Fig. 4), with pitfall traps, including aquatic pitfall traps, and searches yielding more 

hunters than web spinners. All remaining methods yielded more web spinners than hunters.

The importance of using multiple sampling methods to inventory spiders was evidenced by the 

number of species collected by only one method, i.e., method-unique species (Coddington et al.

1996, Standen 2000, Sørensen et al. 2002). Fully 142 (47.0 percent) of the 302 species collected at 

Milbridge were taken exclusively by only one sampling method. Of these method-unique species, 

67 (47.2 percent) were captured by pitfall traps, 33 (23.2 percent) by searches, 28 (19.7 percent) 

by sweep-netting; 6 (4.2 percent) by beating foliage, and 8 (5.6 percent) by litter extraction. Three 

methods, aquatic pitfall traps, shaking foliage over a sweep net, and brushing bark failed to yield 

method-unique species; however, none of these were used extensively during this study. A comparison 

of method-unique species per number of collected specimens showed that litter extractions 

contributed the most (5.4 percent), followed by searches (2.2 percent), pitfall trapping (2.1 percent), 

sweeping (1.8 percent), and beating foliage (1.0 percent).

All inventoried families of spiders at Milbridge were represented by at least one method-unique 

species; two families, Pholcidae and Titanoecidae, were represented solely by method-unique species, 

both collected by individual searches. Overall, more method-unique species of web spinners (N = 

91) were taken than method-unique species of hunters (N = 51). Specimens of Linyphiinae and 

Erigoninae taken by pitfall traps and litter extraction made up most of the method-unique web 

spinners. However, searches, sweep-netting, and beating foliage also yielded unique species of web 

spinners. Pitfall traps, searches, and sweep-netting were the principal means of collecting methodunique hunters, chiefl y species of Salticidae (N = 11), Lycosidae (N = 10), and Thomisidae (N = 7).

Complementarity, i.e., the extent to which two samples or lists complement each other (Colwell and 

Coddington 1994, Sørensen et al. 2002), was also evident among the methods used at Milbridge. Of 

the 302 species inventoried, 179 (59.3 percent of N) were captured by pitfall traps. Searches yielded 

an additional 75 species, bringing the cumulative percentage of species inventoried to 84.1 percent. 

Inclusion of specimens collected by sweep-netting, but not by pitfall-trapping or searches, boosts the 

total to 289 species or 95.7 percent of N. Fully 97.7 percent of the total inventoried spiders can be 

achieved by adding the 6 method-unique species collected by beating coniferous-tree foliage.
163

Our efforts to collect additional species of Dolomedes by deploying aquatic pitfall traps were thwarted. 

The fl oating pitfall traps designed and used by Graham et al. (2003) near a freshwater pond in 

Alberta, Canada, caught numerous specimens of D. triton (Walckenaer 1837), a species likely to 

occur in Milbridge. However, our fl oating pitfalls yielded only specimens of Pirata piraticus (Clerck 

1757), a species frequently caught by terrestrial pitfall traps.

Spider Habitat Associations

As expected, spider species composition varied among the habitats sampled at Milbridge. For 

example, species inhabiting coniferous-tree foliage (i.e., arboreal species) differed signifi cantly 

from those found on the forest fl oor. Segregation and stratifi cation of available habitats is a wellknown phenomenon among spiders (see reviews by Riechert and Gillespie 1986, Uetz 1991). The 

physiognomy or structural features of habitats has an important infl uence on habitat selection, and 

ultimately on spider species composition of habitats (Uetz 1991). Vegetation structure, diversity, and 

architecture all play signifi cant roles in habitat selection and residency by spiders (Greenstone 1984, 

Gunnarsson 1988, 1990; Jennings et al. 1990; Stratton et al. 1979). Consequently, habitat specifi city 

or preferences were indicated by several species of spiders making up the araneofauna of Milbridge.

Species of spiders found in only one habitat (i.e., single-specimen species) might be considered 

habitat-unique species; however, because of their dispersal capabilities (aerial ballooning, cursorial 

activities) such one-specimen species may simply be transients passing through the sampled habitat. 

Hence, caution must be exercised in assigning habitat specifi city or affi nity of single-specimen species.

Species of spiders found in diverse, multiple habitats might be considered habitat-generalist species. 

Nonetheless, some of these occupants of diverse habitats also demonstrated a degree of habitat 

selection or specifi city by their relative abundances. We arbitrarily choose the following criteria as 

indicators of habitat specifi city: 1) species abundance ≥ 10 individuals per habitat, and 2) abundance 

of species X in Y habitat ≥ 66.7 percent, i.e., the proportional distribution of species X abundances 

among all habitats where it was found. Where numerous habitats are sampled, such percentages 

provide an indication of species-habitat specifi city or lack thereof. This was especially so for species 

frequently collected and with high abundances. However, we do not suggest that species meeting 

these criteria are restricted to each specifi ed habitat.

Seashores

Fifty-seven species of spiders were collected in seashore habitats, including cobble, shingle, and grasssedge beaches, and coastal ledges. Of these 57 species, 7 were habitat-unique species, i.e., collected 

exclusively at the seashore: Diplocephalus cristatus (Blackwall 1833), Erigone aletris Crosby & Bishop 

1928, E. zographica Crosby & Bishop 1928, Titanoeca americana Emerton 1888, Pardosa groenlandica 

(Thorell 1872), P. lapidicina Emerton 1885, and Dolomedes striatus Giebel 1869. Most of these 

habitat-unique species were taken by pitfall traps; the lycosids P. groenlandica and P. lapidicina were 

taken chiefl y by searches.

Only 7 of the 57 species of spiders associated with seashores were represented by 10 or more 

individuals. Of these 7 species, 4 showed habitat specifi city ( 66.7 percent of species N) for seashores: 

Diplostyla concolor (Wider 1834), 71.4 percent (N = 42); Diplocephalus cristatus (Blackwall 1833), 

100.0 percent (N = 13); Pardosa groenlandica (Thorell 1872), 100.0 percent (N = 37); and P. 

lapidicina Emerton 1885, 100.0 percent (N = 14). In addition to habitat affi nity, Diplostyla concolor 
164

and Diplocephalus cristatus were taken solely by pitfall traps in a grassy area above the wrack line. 

Pardosa groenlandica and P. lapidicina were found chiefl y on cobble beaches; specimens of both species 

were taken by search and aspirator.

Seashore-Backshores

By sampling the ecotone between seashores and forest or shrub edges, we collected an additional 44 

species of spiders. Of these 44 species, only 2 were habitat-unique species: Erigone ephala Crosby & 

Bishop 1928 and Clubiona moesta Banks 1896. However, both species were represented by single 

specimens. Erigone ephala was swept from backshore grasses; C. moesta was found on alders near the 

shore.

Five of the 44 species associated with seashore-backshores were represented by 10 or more specimens: 

Enoplognatha ovata (Clerck 1757), Grammonota angusta Dondale 1959, Clubiona trivialis C. L. Koch 

1843, Pelegrina fl aviceps (Kaston 1973), and P. fl avipes (Peckham & Peckham 1888). However, none 

of these species showed a preference for the seashore-backshore habitat. Specimens of E. ovata were 

taken chiefl y by sweeping herbaceous-shrub vegetation; the remaining four species were taken most 

frequently by beating coniferous-tree foliage.

Collectively, seashores and seashore-backshores yielded a total of 101 species of spiders; however, only 

9 species were shared in common between these two sampled habitats (QS = 17.8), thus indicating 

that the habitats were distinct. Habitat-unique species in each of the two sampled habitats, and 

arboreal species associated with backshore conifers, no doubt contributed to the observed distinction.

Amelanchier-Rubus Stand

Pitfall-trapping and searches in a stand of serviceberry-blackberry near the seashore yielded 20 

species of spiders. None were unique to this habitat, nor were they captured in suffi cient numbers to 

indicate habitat specifi city. Searches yielded 4 species of web spinners; pitfall traps yielded 5 species 

of web spinners and 11 species of hunters. Two of the hunter species, Zelotes exiguoides Platnick & 

Shadab 1983 and Xysticus ellipticus Turnbull, Dondale & Redner 1965, were method-unique species; 

both were also found in other habitats. Pitfall catches in the Amelanchier-Rubus stand were often 

contaminated by earthworms and slugs, thus making retrieval of spider specimens diffi cult.

Old Fields

Sweep-netting and searches of old fi eld vegetation (grasses, forbs, shrubs, tree seedlings) yielded 75 

species of spiders. Only four were habitat-unique and method-unique species: Ceraticelus bulbosus 

(Emerton 1882), C. emertoni (O. Pickard-Cambridge 1874), Gonatium crassipalpum Bryant 1933, 

and Misumenops carletonicus Dondale & Redner 1976. All four were taken by sweeping old fi eld 

vegetation, but were represented by only one or two specimens. Two specimens of the thomisid M. 

carletonicus were taken from an old fi eld-abandoned farm habitat on Bois Bubert Island; the three 

remaining species (erigonids) were taken from inland old fi elds.

Of the 75 old fi eld species, 13 had abundances ≥ 10; however, none were taken in suffi cient numbers 

to indicate habitat specifi city. The proportional representation of two species, Linyphia triangularis 

(Clerck 1757), 59.0 percent (N = 39) and Xysticus emertoni Keyserling 1880, 54.8 percent (N = 31), 

showed a weak affi liation with old fi eld habitats. Linyphia triangularis is an invasive species frequently 

found in shrubs and small trees of edge habitats.
165

Old Field Edges

Forty-four species of spiders were collected from old fi eld edges, chiefl y by beating coniferoustree foliage and by sweeping herbaceous and shrubbery vegetation. Of the 44 species, only 1 was 

a habitat-unique and method-unique species, i.e., Araneus iviei (Archer 1951) taken by search. 

Fourteen of the 44 old fi eld edge species had abundances ≥ 10 individuals; however, none were taken 

in suffi cient numbers to qualify as specialist of this habitat. Two species, Xysticus punctatus Keyserling 

1880 and Pelegrina fl aviceps (Kaston 1973), showed a weak affi nity for old fi eld edges, specifi cally 

with conifers. The proportional representation of each species’ total collected individuals was X. 

punctatus, 57.1 percent (N = 21); P. fl aviceps, 55.7 percent (N = 70).

Twenty-eight species of spiders were shared in common between old fi elds and old fi eld edges; 

nonetheless, their faunas were distinct, i.e., QS = 47.1.

Old Field-Alder Edge

Twenty-six species of spiders were captured in pitfall traps placed in an old fi eld-alder edge. Of the 

26 species, only 1, Centromerus sylvaticus (Blackwall 1841), was taken exclusively in this habitat and 

exclusively by pitfall traps, thus qualifying it as a habitat-unique and method-unique species. This 

linyphiid was also captured in suffi cient numbers to indicate habitat specifi city; fully 100.0 percent of 

this species’ individuals (N = 19) were trapped in the old fi eld-alder edge.

Of the 26 species collected in this habitat, 11 were shared in common with old fi eld edges; 18 were 

shared in common with old fi elds. However, all three habitats had distinct spider faunas; old fi eldalder edge vs. old fi eld edges, QS = 31.43; old fi eld-alder edge vs. old fi elds, QS = 35.6. Such faunal 

distinctiveness in closely allied habitats was probably due to differences in habitat structure, sampling 

methods, or a combination of both.

Heaths

Seventy-six species of spiders were collected in a Kalmia-Vaccinium heath east of Ficketts Point Road. 

Most were taken by pitfall traps; others were taken by sweeping herbaceous-shrub foliage and by 

litter extraction. Of the 76 species, 5 were unique to this heath: Ceraticelus laticeps (Emerton 1884), 

Metopobactrus prominulus (O. Pickard-Cambridge 1875), Haplodrassus bicornis (Emerton 1909), 

Xysticus ampullatus Turnbull et al. 1965, and X. winnipegensis Turnbull et al. 1965. In addition 

to habitat uniqueness, all fi ve were method-unique species; C. laticeps was taken solely by litter 

extraction and the remainder only by pitfall traps.

Despite the rich diversity of species in this Kalmia-Vaccinium heath, only 7 of the 76 species had 

abundances ≥ 10 individuals, which ranged from 11 to 470. However, only two of the seven 

frequently collected species met or exceeded our defi nition of habitat specifi city: Zelotes exiguoides 

Platnick & Shadab 1983, 86.7 percent (N = 15); and Xysticus winnipegensis Turnbull et al. 1965, 

100.0 percent (N = 11). The lycosid Pardosa distincta (Blackwall 1846) showed a weak affi liation 

with the Kalmia-Vaccinium heath; although 60 individuals were trapped in the heath, this number 

represented only 58.2 percent of the total P. distincta individuals over all sampled habitats (i.e., species 

N = 103).

In addition to the Kalmia-Vaccinium heath, limited searches in a commercial blueberry fi eld yielded 

four species, including one habitat-unique and method-unique species, i.e., Schizocosa communis 
166

(Emerton 1885). This fi eld of common lowbush blueberry (Vaccinium angustifolium Ait.) is located off 

of Old County Road in northern Milbridge. The remaining three species of spiders were: Callilepis pluto

Banks 1896, Drassodes neglectus (Keyserling 1887), and Haplodrassus signifer (C. L. Koch 1839). Only H. 

signifer was shared in common with the Kalmia-Vaccinium heath where extensive sampling was done.

Saltmarshes

Pitfall-trapping, sweep-netting, and individual searches yielded 101 species of spiders in saltmarshes. 

Habitat-unique species in saltmarshes were Allomengea dentisetis (Grube 1861), Dictyna minuta 

Emerton 1888, Emblyna manitoba (Ivie 1947), Castianeira gertschi Kaston 1945, Arctosa rubicunda 

(Keyserling 1877), and Micaria gertschi Barrows & Ivie 1942. The two dictynid species were swept 

from saltmarsh vegetation; the others were captured by pitfall traps placed in saltmarsh litter.

Spider abundances exceeded 10 specimens per species for 21 of the 101 saltmarsh spiders; however, 

only 3 showed defi nite affi nities for saltmarshes. Their proportional representation in saltmarshes 

was Pachygnatha brevis Keyserling 1884, 80.8 percent (N = 26), Arctosa rubicunda (Keyserling 1877), 

100.0 percent (N = 18), and Pardosa modica (Blackwall 1846), 88.0 percent (N = 287). Specimens of 

A. rubicunda were taken exclusively by pitfall traps at Beaver Meadow Brook saltmarsh. Specimens of 

Pardosa modica were also taken at this sampling locality; most were captured by pitfall traps, often in 

large numbers. Fewer specimens of P. modica were taken by searching saltmarsh litter and by sweeping 

saltmarsh vegetation. Sweeping saltmarsh vegetation (Spartina, Juncus) yielded adult specimens of 

Pachygnatha brevis.

An additional three species were collected in suffi cient numbers to show a weak affi liation with 

saltmarshes: Pirata minutus Emerton 1885, 62.5 percent (N = 16), P. piraticus (Clerck 1757), 61.3 

percent (N = 31), and Sergiolus ocellatus (Walckenaer 1837), 62.5 percent (N = 16). All three of these 

cursorial ground-dwellers were taken chiefl y by pitfall traps placed in saltmarsh litter.

Saltmarsh Edges

We sampled saltmarsh edges at both Beaver Meadow Brook and Sawyer Saltmarsh. At Beaver 

Meadow Brook, sweep-netting, searches, and beating coniferous-tree foliage were the principal 

collecting methods. These same methods plus pitfall traps were used to sample the edges of Sawyer 

Saltmarsh. Edges of Cain’s Creek saltmarsh were not sampled.

Collectively, 77 species of spiders were taken from saltmarsh edges. Of these 77 species, 5 were 

habitat-unique species, i.e., taken exclusively in saltmarsh edges. The fi ve habitat-unique species 

were Floricomus rostratus (Emerton 1882), Schizocosa crassipalpata Roewer 1951, Micaria elizabethae 

Gertsch 1942, Misumenops asperatus (Hentz 1847), and Ozyptila sp. (nr. distans Dondale & Redner 

1975). In addition to habitat uniqueness, all fi ve were method-unique species; however, each was 

represented by only one or two individuals.

Saltmarsh edges yielded 11 species of spiders with abundances ≥ 10; only 2 of the 11 were collected 

in suffi cient numbers to indicate habitat specifi city. Fully 68.8 percent of all Dictyna brevitarsa 

Emerton 1915 specimens (N = 16) were taken by beating coniferous-tree foliage along saltmarsh 

edges. Likewise, 68.6 percent of all Arctosa emertoni Gertsch 1934 specimens (N = 70) were captured 

by pitfall traps placed along saltmarsh edges. This ground-dwelling lycosid was also found within 

saltmarsh interiors, but in fewer numbers.
167

Segregation of habitats was indicated by the two species of Arctosa collected during this study. Arctosa 

rubicunda was found solely within the interior of the Beaver Meadow Brook saltmarsh; whereas, A. 

emertoni was found chiefl y along the edges of Sawyer Saltmarsh, and only occasionally (28.6 percent, 

N = 70) within the interior of Beaver Meadow Brook saltmarsh. Additional data (i.e., pitfall traps 

deployed along replicated line transects oriented perpendicular to edges) are needed to confi rm this 

habitat-segregation hypothesis.

As might be expected, 37 species were shared in common between saltmarsh interiors and saltmarsh 

edges. Nonetheless, the two sampled habitats had distinct (QS = 41.6) spider faunas due largely to 

habitat-unique species and relatively few species shared in common.

Salt Meadow

Litter extraction and sweep-netting yielded only 15 species of spiders in a salt meadow near the Bar 

Island causeway; however, our collecting efforts were minimal in this habitat. Two of the 15 species 

were habitat-unique and method-unique: Grammonota maritima Emerton 1925 and Walckenaeria 

spiralis (Emerton 1882). Both were taken by hand-sorting condensed litter. Four of the 15 salt 

meadow species had abundances ≥ 10 individuals; Ceratinops annulipes (Banks 1892), Grammonota 

gigas (Banks 1896), Argiope trifasciata (Forskål 1775), and Ghelna canadensis (Banks 1897). 

Specimens of A. trifasciata were taken by sweeping herbaceous vegetation; specimens of the remaining 

four species were taken by hand-sorting condensed litter. Two of these four frequently collected 

species were taken in suffi cient numbers to indicate habitat specifi city for the salt meadow: Ceratinops 

annulipes, 97.3 percent (N = 37) and G. canadensis, 77.8 percent (N = 18).

Litter samples from the salt meadow yielded numerous juveniles of Ghelna canadensis, a distinctly 

marked species. Besides the adults, the juveniles represented at least three distinct body sizes, possibly 

indicating differential growth rates or multiple generations. Juveniles were not included in the species 

count.

The salt meadow shared few species in common with either saltmarshes or saltmarsh edges; salt 

meadow vs. saltmarshes, QS = 15.5; salt meadow vs. saltmarsh edges, QS = 10.9. However, these low 

similarity percentages may have been due to differences in sampling methodologies and intensity. 

Although samples of litter were taken twice in the salt meadow, pitfall traps that sample continuously 

over longer periods of time were not used in this habitat.

Brackish Marshes

Sweep-netting, searches, and sifted-litter extractions yielded 50 species of spiders in brackish marshes; 

one at the interface of an old fi eld and the Narraguagus River; two further inland, east of Kansas 

Road, and west of Tom Leighton Point Road; and one near a seashore-cobblestone dam south of 

McClellan Park. All were subject to fl ooding during high tides and coastal storms.

Of the 50 brackish-marsh species, 5 were both habitat-unique and method-unique species: Ceratinella 

parvula (Fox 1891), Araneus juniperi (Emerton 1884), Emblyna sublata (Hentz 1850), Clubiona 

pygmaea Banks 1892, and an undetermined genus, species. Ceratinella parvula was taken by siftedlitter extraction; the remaining four species by sweeping brackish marsh vegetation. Seven of the 

50 species associated with brackish marshes had abundances ≥ 10 individuals: Hypselistes fl orens (O. 

Pickard-Cambridge 1875), Tetragnatha laboriosa Hentz 1850, Hypsosinga pygmaea (Sundevall 1831), 
168

Pardosa moesta Banks 1892, Clubiona riparia L. Koch 1866, Phidippus clarus Keyserling 1885, and 

Sitticus fl oricola palustris (Peckham & Peckham 1883). However, none of these seven species were 

taken in suffi cient numbers to indicate a distinct preference for brackish marshes. Most were collected 

by sweeping herbaceous vegetation; H. pygmaea exclusively by sweeping. Dry infl orescences of forbs 

and tree leaves impaled on rushes yielded specimens of S. fl oricola palustris in nesting retreats.

Brackish marshes shared almost twice as many species in common with saltmarshes than with 

freshwater marshes; nonetheless, their associated spider faunas were distinct; brackish marshes vs.

saltmarshes, QS = 43.7; brackish marshes vs. freshwater marshes, QS = 38.3.

Freshwater Marshes

Sweep-netting and searches yielded 44 species of spiders associated with freshwater marshes. Of these 

44 species, four were habitat-unique species: Pardosa fuscula (Thorell 1875), Clubiona maritima L. 

Koch 1867, Marpissa grata (Gertsch 1936), and Synageles noxiosus (Hentz 1850). The same four 

were also method-unique species: P. fuscula and C. maritima by searches; M. grata and S. noxiosus by 

sweeping herbaceous vegetation. All but P. fuscula (N = 5) were represented by single specimens.

Nine of the 44 species associated with freshwater marshes had abundances ≥ 10 individuals; however, 

only 4 exhibited habitat specialty: Tetragnatha caudata Emerton 1884, 88.0 percent (N = 25); 

Clubiona norvegica Strand 1900, 86.7 percent (N = 15); Philodromus peninsulanus Gertsch 1934, 92.9 

percent (N = 14); and Xysticus chippewa Gertsch 1953, 90.0 percent (N = 20). One other species, 

Tibellus maritimus (Menge 1875), exhibited a weak affi nity for freshwater marshes, i.e., 55.6 percent 

(N = 45); however, this species was also taken in saltmarshes, brackish marshes, old fi elds, and other 

habitats.

Although 23 species of spiders were shared in common between freshwater marshes and saltmarshes, 

their inventoried faunas were distinct, i.e., QS = 31.7.

Freshwater Marsh Edges

Fifty-one species of spiders were collected along the edges of freshwater marshes. Most specimens 

were captured by pitfall traps; a few were collected by searching foliage of shrubs. Ten of the 51 

species were both habitat-unique species and method-unique species, i.e., taken exclusively in 

freshwater marsh edges and solely by pitfall traps. The 10 were Robertus banksi (Kaston 1946), 

Bathyphantes gracilis (Blackwall 1841), Centromerus longibulbus (Emerton 1882), Porrhomma terrestre 

(Emerton 1882), Tapinopa bilineata Banks 1893, Carorita limnaea (Crosby & Bishop 1927), 

Walckenaeria redneri Millidge 1983, Antistea brunnea (Emerton 1909), Cybaeopsis euopla (Bishop & 

Crosby 1935), and Scotinella pugnata (Emerton 1890). All but S. pugnata are web spinners.

Spider abundances exceeded or equaled 10 specimens for only 7 of the 51 species associated with 

freshwater marsh edges. Spider-habitat specifi city was indicated by: Bathyphantes pallidus (Banks 

1892), 73.6 percent (N = 110); Antistea brunnea (Emerton 1909), 100.0 percent (N = 27); and Pirata 

insularis Emerton 1885, 84.0 percent (N = 25). All three species were taken by pitfall traps deployed 

in marsh litter.

Searches of dry infl orescences and green leaves of meadowsweet along freshwater marsh edges yielded 

salticid females in nesting retreats. Females of Sitticus fl oricola palustris (Peckham & Peckham 1883) 
169

were fairly common in these microhabitats, sometimes in aggregations of four or fi ve retreats per dry 

infl orescence.

Thirteen species of spiders were shared in common between freshwater marshes and their edges; 

however, the compared faunas were distinct per Sørensen’s similarity quotient, where QS = 27.4. Such 

distinction may be due to differences in strata sampled or sampling methods, or both. Most marsh 

edge specimens came from the ground-litter layer and by pitfall traps; whereas most marsh proper 

specimens came from the herbaceous-shrub layer and by sweeping.

Freshwater Pond

Sweep-netting, searches, and fl oating pitfall traps yielded 27 species of spiders associated with a 

human-made freshwater pond. Only one of the collected species was unique to this habitat, i.e., 

Clubiona obesa Hentz 1847. This species also qualifi ed as a method-unique species because only 

one specimen was found in a retreat on willow. Of the 27 species, none were found in suffi cient 

numbers to indicate a defi nite habitat affi nity. Most of the web spinner species (N = 17) were taken 

by sweeping herbaceous vegetation along the pond shore; species of hunters (N = 10) were taken by 

sweeping, searches, and aquatic pitfall traps.

Fully 70.0 percent of all tetragnathid species inventoried at Milbridge (N = 10) were associated with 

this particular habitat: Pachygnatha brevis Keyserling 1884, Tetragnatha caudata Emerton 1884, T. 

elongata Walckenaer 1842, T. extensa (Linnaeus 1758), T. laboriosa Hentz 1850, T. straminea Emerton 

1884, and T. versicolor Walckenaer 1842. These observations suggest exploitation and species packing 

of an artifi cially made habitat, similar to those described by Wilson (1992).

Freshwater Streamsides

Thirty species of spiders were collected along freshwater streams, including a beaver fl owage near 

Milbridge Elementary School, a dry intermittent stream near Wyman, and Mill River in northwestern 

Milbridge. Most of the specimens were taken by sweeping herbaceous vegetation in these riparian 

habitats; others were taken by searches, beating foliage of conifers, and sifting streamside litter, which 

was processed by Berlese funnels.

None of the 30 freshwater-streamside species were unique to this habitat; only 2 had abundances ≥

10 individuals per species, but neither qualifi ed as a riparian-habitat specialist. Both species, Neoscona 

arabesca (Walckenaer 1842) and Sitticus fl oricola palustris (Peckham & Peckham 1888), were taken in 

diverse habitats at Milbridge. Beating foliage of streamside red spruce yielded specimens of arboreal 

species such as Pityohyphantes phrygianus (C. L. Koch 1836), Grammonota angusta Dondale 1959, and 

Philodromus exilis Banks 1892. Samples of streamside grass litter yielded a few individuals of Pirata 

cantralli Wallace & Exline 1978 and Micaria pulicaria (Sundevall 1831), species found in similar 

habitats such as edges of freshwater ponds and freshwater marshes.

Four of the 30 riparian species, Tetragnatha caudata Emerton 1884, Hypsosinga pygmaea (Sundevall 

1831), Philodromus peninsulanus Gertsch 1934, and Tibellus maritimus (Menge 1875), were methodunique species, i.e., taken solely by sweeping herbaceous vegetation. These four species were also 

found in similar freshwater habitats.
170

Although the freshwater habitats sampled at Milbridge were somewhat similar in vegetative structure 

and composition, their respective spider faunas appeared to be distinct; freshwater streamsides vs.

freshwater pond, QS = 36.4; freshwater streamsides vs. freshwater marshes, QS = 46.0; freshwater 

streamsides vs. freshwater marsh edges, QS = 27.2. However, sampling methods varied among these 

habitats, which may have contributed to the apparent dissimilarities in spider-faunal composition.

Sphagnum Bogs

Two small sphagnum bogs were sampled for spiders; one in west Milbridge, the other near the 

seashore on Bois Bubert Island. Due to their small size and time constraints, neither bog was sampled 

extensively. Only 14 species of spiders were taken in these bogs: 12 species at the inland site, 3 species 

on Bois Bubert Island, with 1 species, Sitticus fl oricola palustris (Peckham & Peckham 1883) shared 

in common. Most of the species were taken by sweeping bog vegetation; three species were taken by 

searches; one species, Walckenaeria directa (O. Pickard-Cambridge 1874), was taken by sifting and 

hand-sorting litter. Both foraging guilds (web spinner, hunter) were represented equally, with seven 

species each.

None of the 14 species were unique to these sphagnum bogs; only 1 species, Argiope trifasciata 

(Forskål 1775), was represented by ≥ 10 individuals. We suspect that additional species of spiders 

inhabit these bogs, especially in the litter layer. Deployment of pitfall traps no doubt would yield 

additional species. The relatively low number of species was probably due to minimal sampling 

efforts.

Roadside Disturbed Areas

Forty-two species were collected in roadside disturbed areas, mostly by sweeping herbaceous 

vegetation along drainage ditches and rights-of-way. A few specimens were taken by searching under 

rocks and inside culverts; fewer still by beating roadside shrubs and trees. None of the 42 species 

were unique to roadsides; only 1 species, Enoplognatha ovata (Clerck 1757), was represented by ≥ 10 

individuals, but not in suffi cient numbers to demonstrate a habitat specifi city for roadsides. In fact, 

this theridiid spider was found in diverse habitats at Milbridge and usually associated with herbaceous 

vegetation.

Not surprisingly, species of web spinners were more prevalent among these roadside collections 

than species of hunters, due chiefl y to our reliance on sweep-netting for sampling these habitats. In 

addition to Enoplognatha ovata, two other invasive species were found in roadside disturbed areas. 

The theridiid Achaearanea tabulata Levi 1980 was found in roadside culverts; the linyphiid Linyphia 

triangularis (Clerck 1757) was found on roadside Juncus sp.

Betula-Acer Stand

Pitfall-trapping in forest-fl oor litter, sweeping understory vegetation, brushing bark, and searches 

yielded 27 species of spiders associated with a mixed birch-maple stand near Narraguagus Bay. Most 

of the 27 species were taken by pitfall traps; only 2 species were swept from understory vegetation. 

Only one species was taken exclusively in this habitat, i.e., Enoplognatha intrepida (Sørensen 1898) 

found under the bark of white (paper) birch, thus qualifying as both a habitat-unique and methodunique species. Two of the 27 associated species had collection abundances ≥ 10 individuals per 

species: Agelenopsis utahana (Chamberlin & Ivie 1933) and Neoantistea magna (Keyserling 1887). The 

latter species showed a distinct affi nity for this forest-stand type. Fully 233 specimens, representing 
171

82.3 percent of all collected individuals of N. magna, were captured by pitfall traps in this birchmaple stand. Although N. magna was found in other sampled habitats, none of the observed 

abundances approached that of the Betula-Acer stand.

Bigtooth Aspen Stand

Twenty-nine species were collected in a mature bigtooth aspen stand near Sawyer Saltmarsh. All 29 

species were captured by pitfall traps. None were habitat-unique species, and only 4 had abundances 

≥ 10 individuals per species: Pardosa xerampelina (Keyserling 1877), Phrurotimpus alarius (Hentz 

1847), Zelotes fratris Chamberlin 1920, and Xysticus elegans Keyserling 1880. Not surprisingly, with 

such low abundances, none of these four species showed a distinct affi nity for the ground-litter layer 

of this bigtooth aspen stand; habitat-species percentages ranged from 7.6 percent to 18.2 percent.

Aspen-Maple Stand

Condensed litter samples extracted by Berlese funnel yielded fi ve species associated with the 

forest fl oor in an aspen-maple stand of east Milbridge. One of the fi ve was a habitat-unique and 

method-unique species, but it was represented by only one specimen: Eperigone undulata (Emerton 

1914). None of the fi ve species associated with this habitat were collected in suffi cient numbers to 

demonstrate habitat specifi city.

White Birch Stand

Pitfall-trapping and sweeping understory vegetation yielded 39 species of spiders associated with 

a mature white birch stand near Sawyer Saltmarsh. Of the 39 species, none were habitat-unique; 

however, 3 were method-unique species captured by pitfall traps. Five specimens of Mangora placida 

(Hentz 1847) were swept from understory vegetation in this stand; the remaining 38 species were 

taken by pitfall traps.

Nine of the 39 species taken in the white birch stand had abundances ≥ 10 individuals; however, none 

were taken in suffi cient numbers to demonstrate habitat specifi city. Pitfall-trap catches of Amaurobius 

borealis Emerton 1909 showed a weak habitat affi liation, with 56.4 percent of all individuals of 

this species taken in the white birch stand. Xysticus elegans Keyserling 1880 was the most frequently 

captured species; however, it was also captured by pitfall traps in other deciduous-tree stands.

Red Maple-White Birch Stand

Thirty-eight species of spiders were associated with a mature red maple-white birch stand west of 

Sawyer Saltmarsh. All were captured by pitfall traps placed in forest-fl oor litter. None of the 38 

species were habitat-unique, but 2 were method-unique species: Amaurobius borealis Emerton 1909 

and Agroeca ornata Banks 1892. Only 7 of the 38 species had abundances ≥ 10 individuals. In 

addition to Amaurobius borealis and Agroeca ornata, frequently captured species in this habitat were: 

Callobius bennetti (Blackwall 1846), Alopecosa aculeata (Clerck 1757), Phrurotimpus alarius (Hentz 

1847), Zelotes fratris Chamberlin 1920, and Xysticus elegans Keyserling 1880. However, none were 

taken in suffi cient numbers to indicate a preference for this red maple-white birch stand.

Coastal Red Spruce Stands

Pitfall traps and litter-extracted samples yielded 31 species of spiders associated with two stands of 

coastal red spruce: one at McClellan Park where pitfall traps were used, and one on Bar Island where 

samples of forest-fl oor litter were collected and spiders extracted by Berlese funnels. Of the 31 species 

of spiders associated with these stands, 3 were both habitat-unique and method-unique species; 4 
172

specimens of Lepthyphantes alpinus (Emerton 1882) captured by pitfall traps; 1 specimen each of 

Eperigone entomologica (Emerton 1911) and Sciastes truncatus (Emerton 1882) by litter-extracted 

samples.

Only 3 of the 31 species of spiders associated with red spruce stands had abundances ≥ 10 

individuals, all taken by pitfall traps. The three species captured in abundance were Cryphoeca 

montana Emerton 1909, Neoantistea magna (Keyserling 1887), and Callobius bennetti (Blackwall 

1846). However, only Cryphoeca montana was taken in suffi cient numbers to demonstrate a distinct 

habitat affi nity for forest-fl oor litter of red spruce; fully 92.1 percent of all C. montana specimens 

(N = 76) came from this habitat. Although 35 specimens of N. magna were also captured, their 

proportional representation in the red spruce stand was minimal (12.4 percent) compared to a 

Betula-Acer stand where 233 specimens (82.3 percent) were captured by pitfall traps. These results 

indicate that N. magna showed a distinct habitat specifi city for birch-maple stands over red spruce 

stands at Milbridge. This hahniid was also a method-unique species; it was taken solely by pitfall traps 

regardless of forest-stand type.

Mixed Conifer Stands

Mixed conifer stands, chiefl y red spruce and balsam fi r, were sampled for spiders at several inland and 

offshore sites in Milbridge. Fifty species of spiders were taken from diverse microhabitats in mixedconifer stands. Most were collected by beating foliage of conifers; others were taken by sweeping 

understory vegetation (Kalmia, Spiraea) and by searching forest-fl oor litter, ledges, and under rocks 

and loose tree bark. Of the 50 species, 2 were both habitat-unique and method-unique species: 

Theridula emertoni Levi 1954 and Estrandia grandaeva (Keyserling 1886). Both species were taken on 

Bois Bubert Island, where T. emertoni was swept from Kalmia sp. and E. grandaeva was beaten from 

foliage of red spruce.

Of the 50 species of spiders associated with mixed conifer stands, only Callobius bennetti (Blackwall 

1846) had abundances ≥ 10 individuals, but insuffi cient numbers to demonstrate habitat specifi city. 

This amaurobiid spider was found in diverse habitats at Milbridge: chiefl y under loose bark of 

standing and downed trees, but also under rocks and in crevices of ledges.

The spider fauna associated with mixed conifer stands was allied to that found in mixed coniferhardwood stands; QS = 52.5. Fully 26 species were shared in common between these two forest-stand 

types. However, the spider fauna associated with mixed hardwood-conifer stands apparently differs 

(QS = 37.8) from that found in mixed conifer stands. By defi nition, QS values less than 50.0 are 

considered distinct (Price 1975).

Mixed Conifer-Hardwood Stands

Forty-nine species of spiders were collected from diverse habitats at several inland and offshore sites. 

Most of the species were taken by beating foliage of conifers and sweeping understory vegetation. A 

few were found under rocks, on the ground, and beneath loose bark of hardwoods. Only 1 of the 49 

associated species was unique to this forest-stand type; a single specimen of Dismodicus decemoculatus 

(Emerton 1911) was swept from laurel on Bois Bubert Island, thus making it a method-unique 

species at Milbridge.
173

The inventoried spider fauna associated with mixed conifer-hardwood stands differed from that 

associated with mixed hardwood-conifer stands (QS = 38.9), which had several habitat-unique 

species.

Mixed Hardwood-Conifer Stands

Sixty-four species of spiders were collected in mixed hardwood-conifer stands at Milbridge. All 

sampled sites were inland. Most of the species were taken by pitfall traps placed in a hardwoodconifer woodland near the eastern shore of Narraguagus Bay. Others were collected by sweeping 

understory vegetation, searching ground litter, ledges, and foliage of small trees, and beating foliage of 

hardwoods and conifers. A few species were collected by sifting and hand-sorting litter beneath maple 

and birch trees.

Of the 64 species, 6 were both habitat-unique and method-unique species: Rugathodes aurantius 

(Emerton 1915), Meioneta nigripes (Simon), Bathyphantes brevis (Emerton 1911), Hypsosinga rubens 

(Hentz 1847), Cicurina arcuata Keyserling 1887, and C. placida Banks 1892. All except H. rubens

were captured by pitfall traps and were represented by only one or two specimens. A single female of 

H. rubens was taken by beating foliage of balsam fi r.

Only 3 of the 64 species were represented by ≥ 10 individuals; however, species abundances were 

not suffi cient to demonstrate habitat specifi city for this forest-stand type. Although tree species were 

shared in common between mixed conifer-hardwood and mixed hardwood-conifer stands, their 

respective spider faunas apparently differed, QS = 38.9. This apparent distinctiveness may have been 

due to differences in sampling methodologies. Although most of the same methods were used in both 

stand types, pitfall traps were not used in mixed conifer-hardwood stands. Only one habitat-unique 

species was found in the mixed conifer-hardwood stands compared to six in the mixed hardwoodconifer stands, all but one taken by pitfall traps.

Gravel Pit

Only 11 species of spiders were collected in a gravel pit northeast of Milbridge village. The gravel pit 

is located in a conifer-mixed hardwood stand. Specimens were taken chiefl y by pitfall traps; a few 

were taken by searches. None of the 11 species were unique to this habitat, nor were they collected 

in suffi cient numbers to indicate habitat specifi city. However, four species, Diplostyla concolor (Wider 

1834), Castianeira descripta (Hentz 1847), Habronattus viridipes (Hentz 1846), and Phidippus 

purpuratus Keyserling 1885, were method-unique species; the fi rst three were taken exclusively by 

pitfall traps, the latter solely by searches. Of the 11 collected species, 4 were web spinners; 7 were 

hunters.

Eight females of Grammonota angusta Dondale 1959, a species found chiefl y on coniferous-tree 

foliage, were captured by pitfall traps placed on the ground in the gravel pit.

School Playground-Disturbed Area

A few sweeping samples were taken in a disturbed area adjacent to the Milbridge Elementary School 

playground. Vegetation of this area resembles that of old fi elds, i.e., grasses and forbs of pioneering 

species. Seventeen species of spiders were collected in this disturbed area; two were both habitatunique and method-unique species: Acanthepeira stellata (Walckenaer 1805) and Eustala cepina 

(Walckenaer 1842). None of the 17 species associated with this habitat were collected in suffi cient 
174

numbers to demonstrate habitat specifi city. Individuals of Evarcha hoyi (Peckham & Peckham 1883), 

Pelegrina insignis (Banks 1892), and Phidippus clarus Keyserling 1885 were relatively abundant among 

the sweepings, but none had collection frequencies ≥ 10 individuals.

Domestic Habitats

Ninety-eight species of spiders were collected exclusively by searches of domestic habitats. These 

habitats included interiors and exteriors of houses, storage shed, barn, screened house, seashore 

cottage, and professional building. Also included in the domestic category were boat, canoe, patio, 

hot tub, lawns and lawn furniture, ornamental shrubs and trees, compost bins, and vegetable and 

fl ower gardens.

Of the 98 species found in domestic habitats, most can be considered as “accidentals” or transient 

invaders. For example, 15 of the 98 species qualifi ed as habitat-unique species, but few are known 

to be synanthropic species. Most of these “accidentals” were represented by only one or two 

specimens. The domestic habitat-unique species were Pholcus manueli Gertsch 1937, Steatoda 

americana (Emerton 1882), S. borealis (Hentz 1850), Takayus lyricus (Walckenaer, 1842), Meioneta 

fabra (Keyserling 1886), Soulgas corticarius (Emerton 1909), Tapinocyba sp. (unknown), Meta ovalis 

(Gertsch 1933), Araneus cavaticus (Keyserling 1882), Pardosa saxatilis (Hentz 1844), Herpyllus 

ecclesiasticus Hentz 1832, Philodromus praelustris Keyserling 1880, P. vulgaris (Hentz 1847), Euophrys 

monadnock Emerton 1891, and Sitticus pubescens (Fabricius 1775). Only one of these habitat-unique 

species can be considered domestic sensu stricto, the invasive hunter S. pubescens. The web spinners 

Pholcus manueli and A. cavaticus frequent domestic habitats, but can be found elsewhere in natural 

habitats. No doubt the constancy of observer searches contributed to the apparent spider-faunal 

richness of domestic habitats.

Invasive Species

At least 12 of the 302 species of spiders collected at Milbridge were non-native invasives, chiefl y from 

the Palearctic realm; these include Achaearanea tabulata Levi 1980, Enoplognatha ovata (Clerck 1757), 

Neottiura bimaculata (Linnaeus 1767), Steatoda bipunctata (Linnaeus 1758), Linyphia triangularis 

(Clerck 1757), Pityohyphantes phrygianus (C. L. Koch 1836), Porrhoma convexum (Westring 1851), 

Zygiella atrica (C. L. Koch 1845), Tegenaria domestica (Clerck 1757), Trochosa ruricola (De Geer 

1778), Salticus scenicus (Clerck 1757), and Sitticus pubescens (Fabricius 1775).

Not surprisingly, about half of the invasive species at Milbridge were found in or near domestic 

habitats: Achaearanea tabulata, Steatoda bipunctata, Tegenaria domestica, Salticus scenicus, and Sitticus 

pubescens. Due to their close association with human habitations, these and other synanthropic 

species are prone to widespread, long-range dispersal (Gertsch 1979, Kaston 1983), perhaps some 

dating back to early colonial times or even earlier on the northeastern Atlantic seaboard.

However, the dates, origins, and focal points of such introductions are generally unknown. The 

comb-footed spider A. tabulata was described from a single female taken in August 1976 at 

Richmond Hill, NY (Levi 1980a). This species has since spread rapidly into other regions of New 

England, including Maine and Canada (Dondale et al. 1994). Because of its close similarity to 

Parasteatoda tepidariorum (C. L. Koch), another invasive species fi rst introduced into the Southern 

United States (Gertsch 1979), A. tabulata may have been overlooked earlier in the northeastern 

region (Levi 1980a). Nyffeler et al. (1986) provided evidence that Steatoda bipunctata, an invasive 
175

from Europe, has partially displaced the native S. borealis (Hentz) in the Northeastern United States 

and Southeastern Canada. The apparent sparsity of S. borealis and relative abundance of S. bipunctata 

at Milbridge supports displacement theory; nonetheless, historical data are lacking. The William 

Procter collection of spiders on Mount Desert Island during the l920s and early 1930s provides the 

earliest known records of S. bipunctata in Maine (Nyffeler et al. 1986, Procter 1946).

According to Gertsch (1979), the European Zygiella atrica was fi rst noticed on wharves in 

Massachusetts in about 1880; by 1911, it was abundant along coastal regions of Massachusetts and 

Rhode Island. Zygiella atrica now occurs in maritime habitats along the Maine coast including those 

of Milbridge. The European Sitticus pubescens (Fabricius 1775) was fi rst collected in Allston, MA, the 

earliest record for this species in North America (Bryant 1941). This salticid spider has since been 

found in Connecticut (Kaston 1981) and in New Jersey (Cutler, pers. comm.).

The sheet-web weaver L. triangularis remains to be found elsewhere in New England or in North 

America (Jennings et al. 2002). Nonetheless, this Palearctic species is apt to spread rapidly because of 

its reproductive potential, aerial dispersal capabilities, and aggressive behaviors toward other spiders. 

We suspect that L. triangularis poses a threat to the native araneofauna at Milbridge and elsewhere 

in Maine (Jennings et al. 2002). Unlike some other invasive species, L. triangularis generally inhabits 

herb-shrub-small tree strata away from buildings; however, it also occurs in edge habitats (hedges, 

shrubs, trees) near houses.

Other invasive spiders at Milbridge (i.e., Enoplognatha ovata, Neottitura bimaculata, Pityohyphantes 

phrygianus, and Trochosa ruricola) were generally found away from domestic habitats. The theridiid 

E. ovata lives in deciduous shrubs and trees; it was especially abundant in sweepings of herbaceous 

vegetation (jewelweed) and shrubs along roadsides and forest edges. Neottiura bimaculata occupies the 

herb-shrub layer of old fi elds, marshes, and bogs; at Milbridge we found this species in fi ve habitats, 

only one semidomestic (i.e., overturned canoe in old fi eld). Sheet webs of P. phrygianus were usually 

observed and specimens taken on coniferous-tree foliage (e.g., red spruce), especially along old fi eld 

edges. The lycosid T. ruricola was most abundant near the seashore of Narraguagus Bay. This latter 

species may pose a threat to our native Trochosa terricola Thorell 1856, especially on offshore islands 

where invasives are usually most successful at displacing natives (Cox 1999, Wilson 1992).

The zebra spider, Salticus scenicus (Clerck 1757), is common to both Europe and America and 

inhabits both domestic and native habitats. Although found chiefl y in domestic and semidomestic 

habitats at Milbridge, S. scenicus has been observed inhabiting seashore ledges in Maine (Jennings, 

unpubl.).

Species-Range Extensions

The Milbridge collections yielded signifi cant geographic range extensions for several species of 

spiders, including 31 new state records for Maine, 6 New England records, 3 national records, and 

1 North American record, i.e., Linyphia triangularis (Clerck, 1757), a Palearctic species reported 

earlier (Jennings et al. 2002). Both native and non-native species were included among the observed 

range extensions. For example, before the Milbridge study, Meioneta nigripes (Simon 1884) had been 

found in Québec, the Northwest Territories, Greenland, and the Palearctic (Buckle et al. 2001), but 

not in Maine or elsewhere in the Northeastern United States. Conversely, Erigone blaesa Crosby and 

Bishop, 1928 has been taken in several New England States and northeastern Canadian Provinces, 
176

but not in Maine. The European Erigone dentipalpis (Wider, 1834) found in Great Britain (Locket 

and Millidge 1953, Roberts 1987) has since been introduced (?) into Newfoundland (Buckle et al.

2001). Our collection of E. dentipalpis in Milbridge represents the second locality for this species in 

North America. First described from specimens taken along the seashore in Barrington, NS (Emerton 

1925), and since in Québec and Newfoundland (Buckle et al. 2001), our collections of Grammonota 

maritima Emerton, 1925 from a salt meadow in Milbridge represent the fi rst records of this species in 

Maine, New England, and the Eastern United States. Although previously recorded from Wisconsin, 

Washington, Québec, Manitoba, and Saskatchewan, the collection of Walckenaeria redneri Millidge 

1983 from a freshwater marsh in Milbridge extends the known range of this species to Maine and 

New England. Likewise, specimens of Emblyna manitoba (Ivie, 1947) taken in Milbridge extend the 

range of a species previously known from the upper midwest, the Pacifi c Northwest, and Florida 

(Chamberlin and Gertsch 1958) to Maine and New England. This dictynid spider is also known 

in Ontario, Québec, and some western provinces (Buckle et al. 2001). Our collections of another 

widespread species, Xysticus luctuosus (Blackwall, 1836), extend its previously known range to Maine 

and New England. This crab spider ranges from Alaska, British Columbia, Oregon, Utah, Wyoming, 

Alberta, and Minnesota to Manitoba, Ontario, Québec, New Brunswick, and Labrador (Dondale 

and Redner 1978b; Gertsch 1939, 1953; Turnbull et al. 1965). Our collections of Sibianor aemulus 

(Gertsch, 1934) represent new State and Eastern United States records; before the Milbridge study, S. 

aemulus was known from Alberta, Manitoba, Ontario, Québec, and New Brunswick (Bélanger and 

Hutchinson 1992, Maddison 1978, Richman and Cutler 1978). Before our study, the invasive Sitticus 

pubescens (Fabricius 1775) had been taken only in Connecticut and Massachusetts (Kaston 1981); the 

Milbridge record represents a northward expansion. Apparently this species remains to be found in 

Québec (Paquin and Dupérré 2003).

Faunal Similarities

Predictably, the Milbridge fauna showed greater degrees of similarity among spider families and genera 

than among species when compared to the Mount Desert Island (MDI) fauna; for families, overall 

QS = 87.8; for genera, overall QS = 74.8; for species, overall QS = 52.8. However, the calculated QS 

value for species was only slightly above the arbitrary 50 percent limit for distinguishing similar vs.

distinct communities (Price 1975). Because Milbridge and MDI are within the same East Coastal 

BioPhysical Region, a region with similar biotic and physical features (McMahon 1990), we had 

expected a much higher degree of similarity in spider species composition for the two compared 

faunas. Overall, 1 family, 47 genera, and 173 species were found at Milbridge but not at MDI; 

conversely, 4 families, 12 genera and 49 species were found at MDI but not at Milbridge. Similar 

differences in composition of spider taxa were also evident at the foraging guild level. For web 

spinners, 1 family, 36 genera, and 100 species were found at Milbridge but not at MDI; conversely, 

2 families, 7 genera, and 29 species were found at MDI but not at Milbridge. For hunters, 0 families 

11 genera, and 73 species were collected at Milbridge but not at MDI; whereas 2 families, 5 genera, 

and 20 species were collected at MDI but not at Milbridge. Our results indicated that web spinner 

abundances contributed most to the disparity in similarity of the two compared faunas.

We suspect that differences in sampling methodologies accounted for the relatively low degree of 

species similarity between the two faunas. At Milbridge, we relied chiefl y on pitfall traps to sample 

the ground-inhabiting fauna; whereas at MDI, Procter and his associates used searches (e.g., turning 

stones, logs) and litter sifting to sample the ground-inhabiting fauna. The pitfall traps at Milbridge 

yielded a rich diversity of species not previously recorded as components of the MDI araneofauna. 
177

Chief among these were species of linyphiid spiders commonly associated with ground litter. Notably 

absent from the MDI fauna listed by Procter (1946) were species of Centromerus, Eperigone, Erigone, 

and Walckenaeria, all small web spinners commonly taken by pitfall traps. Also absent from the 

inventoried MDI fauna were ground-dwelling species Arctosa, Schizocosa, and Sergiolus, all cursorial 

hunters commonly captured in pitfall traps. Likewise, additional species of Pardosa, Pirata, Clubiona, 

and Xysticus most likely occur at MDI. For example, the most frequently collected species of spider at 

Milbridge, Pardosa moesta Banks, 1892, surprisingly was not included among the inventoried species 

at MDI (Procter 1946).

Notably absent from the inventoried spiders at Milbridge, but present at MDI, were species of Hyptiotes, 

Theridiosoma, Helophora, Maso, Ero, Mimetus, Elaver, and Misumenoides. Further sampling of the 

fauna at Milbridge should yield additional species of Robertus, Theridion, Neriene, Tetragnatha, Araneus, 

Zygiella, Emblyna, Dolomedes, Drassyllus, Philodromus, Xysticus, and Tutelina, all components of the MDI 

fauna. The invasive Dysdera crocata C. L. Koch, 1838 present at MDI may also occur in Milbridge.

ESTIMATED FAUNAL RICHNESS

Estimated Milbridge Fauna

Like most faunal inventories, the absolute limit of spider species composition was not attained during 

this study. Consequently, we prepared a list of spider families, genera, and species likely to occur in 

Milbridge (Table 2). The list is based on collections of spiders from the same East Coastal BioPhysical 

Region, i.e., chiefl y records from nearby localities in Washington and Hancock Counties. Both 

published and unpublished data sources were used to compile the list.

The additional spider taxa likely to occur in Milbridge include 5 families (Uloboridae, 

Theridiosomatidae, Dysderidae, Mimetidae, and Anyphaenidae), 27 genera (e.g., Hyptiotes, Theonoe, 

Theridiosoma, Helophora), and 109 species (Table 2). Hence, the total estimated araneofauna at 

Milbridge most likely comprises 24 families, 172 genera, and ~ 411 species. Results of this study 

indicate that 79.2 percent of the possible families, 84.3 percent of the possible genera, and 73.5 

percent of the possible species have been found in Milbridge thus far. Of the 109 likely species, fully 

51.4 percent had been found in commercial lowbush blueberry fi elds, which suggests that future 

inventories should concentrate sampling efforts in these habitats. Additional habitats and strata not 

sampled during the current study (e.g., stands of northern white-cedar, mid- and upper-crown levels 

of trees, bog litter) no doubt will lead to new discoveries.

SPIDER-HABITAT CONSERVATION

Spider Conservation

In addition to prey abundance, spiders respond to structural features within the habitat; i.e., physical 

features of the habitat are important determinants of spider species composition and abundances 

(see reviews by Greenstone 1984, Riechert and Gillespie 1986, Uetz 1991). For example, plant 

architecture or physiognomy greatly infl uences both the composition and abundance of spider 

occupants (Gunnarsson 1988, Jennings et al. 1990, Stratton et al. 1979). Because of spiders’ close 

ties with habitat, alterations of habitat (e.g., deforestation, agriculture, grazing, and urbanization) 

can have dire consequences. For example, forest clearcutting drastically reduces spider abundance 

and species composition (Coyle 1981, McIver et al. 1990, 1992); strip clearcutting also affects spider 

abundance and composition (Jennings et al. 1988).
178

Spider taxa 1 Locality record(s) 2 Habitat(s) Source(s)

WEB SPINNERS

PHOLCIDAE

Pholcus phalangioides (Fuesslin, 1775) Machias, Washington County inside house, barn Jennings (unpubl.)

ULOBORIDAE

Hyptiotes cavatus (Hentz, 1847) Mount Desert Island (MDI), in underbrush Procter (1946) 3

 Hancock County

Hyptiotes gertschi Chamberlin & Ivie, 1935 Steuben, Washington County in web, dead spruce limb HWRI (1998 ) 4

THERIDIIDAE

Crustulina altera Gertsch & Archer, 1942 MDI, Hancock County under stones Procter (1946) 3

Enoplognatha caricis (Fickert, 1876) Columbia, Washington County litter, blueberry fi eld Maloney (2002)

MDI, Hancock County (n. d.) 5 Procter (1946) 3

Enoplognatha marmorata (Hentz, 1850) Cherryfi eld, Columbia, Deblois, 

Jonesboro, & T19 MD BPP, Washington County

litter, blueberry fi eld Collins et al. (1996)

Cherryfi eld, Washington County litter, blueberry fi eld Timms (unpubl.)

Cherryfi eld, Columbia, & Jonesboro, Washington County

litter, blueberry fi eld Maloney (2002)

Columbia, T18 MD BPP, & T19 MD BPP, Washington County

litter, blueberry fi eld Drummond et al. (unpubl.)

Euryopis argentea Emerton, 1882 Cherryfi eld, Washington County litter, blueberry fi eld Timms (unpubl.)

Petit Manan National Wildlife Refuge, (PMNWR), Steuben, Washington County

litter, maritime slope bog Jennings (unpubl.)

Cherryfi eld & Jonesboro, Washington County

Maloney (2002)

T18 MD BPP, Washington County litter, blueberry fi eld Drummond et al. (unpubl.)

Euryopis saukea Levi, 1951 T19 MD BPP, Washington County litter, blueberry fi eld Collins et al. (1996)Keijia tincta (Walckenaer, 1802) ANP headquarters, Bar Harbor, MDI, beating white spruce foliage Jennings (unpubl.)

Parasteatoda tepidariorum (C. L. Koch, 1841) MDI, Hancock County in or near houses, buildings; 

cellars

Procter (1946) 3

Acadia Nat’l Park (ANP), Bar Harbor, MDI, Hancock County

guest housing, porch foyer Jennings & Cutler 

(unpubl.)

Table 2.—Spider taxa likely to occur in Milbridge based on published and unpublished records of species elsewhere in the East Coastal BioPhysical Region.

continued
179

Rhomphaea fi ctilium (Hentz, 1850) MDI, Hancock County (n. d.) 5 Procter (1946) 3Robertus borealis (Kaston, 1946) PMNWR, Steuben, Washington 

County

litter, Sphagnum bog Jennings (unpubl.)

Robertus pumilus (Emerton, 1909) Winter Harbor, Hancock County sifted Sphagnum, brackish marsh HWRI (1998) 4

Robertus riparius (Keyserling, 1886) MDI, Hancock County sifting (litter) Procter (1946) 3

Steuben, Washington County sifted leaf litter HWRI (1998) 4

Robertus spinifer (Emerton, 1909) Cherryfi eld & Columbia, Washington 

County

litter, blueberry fi eld Collins et al. (1996)

Cherryfi eld, Washington County litter, blueberry fi eld Timms (unpubl.)

PMNWR, Steuben, Washington County

litter, old fi eld-abandoned pasture Jennings (unpubl.)

Cherryfi eld, Columbia, & Jonesboro, Washington County

litter, blueberry fi eld Maloney (2002)

Columbia, T18 MD BPP, T19 MD BPP, Washington County

litter, blueberry fi eld Drummond et al. (unpubl.)

 Theonoe stridula Crosby, 1906 ANP, Schoodic Peninsula, Winter 

Harbor, Hancock County

sifted litter; hanging bog HWRI (1999) 4 

Theridion petraeum L. Koch, 1872 MDI, Hancock County Sphagnum bog Procter (1946) 3Theridion varians Hahn, 1831 Winter Harbor, Hancock County on spruce, on cedar Houser (unpubl.) 

Thymoites marxi (Crosby, 1906) Steuben, Washington County sweeping roadside vegetation HWRI (1991) 4

THERIDIOSOMATIDAE

Theridiosoma gemmosum (L. Koch, 1877) MDI, Hancock County low, wet places Procter (1946) 3

LINYPHIIDAE (Linyphiinae)

Centromerus denticulatus (Emerton, 1909) Winter Harbor, Hancock County sifted litter; hanging bog HWRI (1999) 4Helophora insignis (Blackwall, 1841) MDI, Hancock County (n. d.) 5 Procter (1946) 3

Winter Harbor, Hancock County beating red spruce foliage HWRI (1999) 4

Megalepthyphantes nebulosus (Sundevall, 1830) MDI, Hancock County (n. d.) 5 Procter (1946) 3Meioneta angulata (Emerton, 1882) MDI, Hancock County sifting (litter) Procter (1946) 3Neriene variabilis (Banks, 1892) MDI, Hancock County (n. d.) 5 Procter (1946) 3Oreonetides sp. 1 Jonesboro, Washington County litter, blueberry fi eld Collins et al. (1996)

Pityohyphantes limitaneus (Emerton, 1915) MDI, Hancock County on bush Procter (1946) 3

Table 2.—continuedSpider taxa 1 Locality record(s) 2 Habitat(s) Source(s)

continued
180

Stemonyphantes blauveltae Gertsch, 1951 Cherryfi eld, Washington County litter, blueberry fi eld Maloney (2002)

Tennesseellum formica (Emerton, 1882) Harrington, Washington County Sphagnum bog HWRI (1997) 4

Columbia & T18 MD BPP, Washington County

litter, blueberry fi eld Drummond et al. (unpubl.)

LINYPHIIDAE (Erigoninae)

Baryphyma longitarsum (Emerton, 1882) Jonesboro, Washington County litter, blueberry fi eld Collins et al. (1996)

Ceratinopsis auriculata Emerton, 1909 Amherst, Hancock County litter, blueberry fi eld Drummond et al. (unpubl.)

 Ceratinopsis interpres (O. Pickard-Cambridge, 1874)

MDI, Hancock County heath Procter (1946) 3

Coloncus sp. (undetermined) Columbia litter, blueberry fi eld Drummond et al. (unpubl.)

Eperigone trilobata (Emerton, 1882) Steuben, Washington County litter, old fi eld-abandoned pasture Jennings (unpubl.)

Cherryfi eld & Jonesboro, Washington County

litter, blueberry fi eld Maloney (2002)

Columbia, Washington County litter, blueberry fi eld Drummond et al. (unpubl.)T18 MD BPP & T19 MD BPP, Washington County

litter, blueberry fi eld Drummond et al. (unpubl.)

Eridantes erigonoides (Emerton, 1882) Cherryfi eld, Washington County litter, blueberry fi eld Maloney (2002)

T18 MD BPP, Washington County litter, blueberry fi eld Drummond et al. (unpubl.)

Erigone autumnalis (Emerton, 1882) T18 MD BPP, Washington County litter, blueberry fi eld Drummond et al. (unpubl.)Floricomus plumalis (Crosby, 1905) Steuben, Washington County sifted litter; deciduous-mixed 

conifer

HWRI (1998) 4 

T18 MD BPP, Washington County litter, blueberry fi eld Drummond et al. (unpubl.)

Grammonota capitata Emerton, 1924 Amherst, Hancock County litter, blueberry fi eld Drummond et al. (unpubl.)

Bar Harbor, Hancock County beating white spruce foliage Jennings (unpubl.)Jonesboro, Washington County litter, blueberry fi eld Collins et al. (1996)Cherryfi eld, Washington County litter, blueberry fi eld Timms (unpubl.)

PMNWR, Steuben, Washington County

litter, mountain ash woodland Jennings (unpubl.)

Cherryfi eld, Columbia, Harrington, & Jonesboro, Washington County

litter, blueberry fi eld Maloney (2002)

Table 2.—continuedSpider taxa 1 Locality record(s) 2 Habitat(s) Source(s)

continued
181

Cherryfi eld, Columbia, T18 MD BPP, T19 MD BPP, & Whitneyville, Washington County

litter, blueberry fi eld Drummond et al. (unpubl.)

Grammonota ornata (O. Pickard- Cambridge,

 1875)

T18 MD BPP litter, blueberry fi eld Drummond et al. (unpubl.)

Grammonota pallipes Banks, 1895 Columbia, Washington County litter, blueberry fi eld Drummond et al. (unpubl.)

Grammonota pictilis (O. Pickard-Cambridge,

 1875)

MDI, Hancock County in woods, on litter, on tree trunk Procter (1946) 3

Idionella rugosa (Crosby, 1905) Cherryfi eld, Washington County litter, blueberry fi eld Maloney (2002)

Lophomma depressum (Emerton, 1882) T18 MD BPP, Washington County litter, blueberry fi eld Drummond et al. (unpubl.)

Maso sundevalli (Westring, 1851) MDI, Hancock County sifting (litter) Procter (1946) 3

Phlattothrata fl agellata (Emerton, 1911) MDI, Hancock County (n. d.) 5 Procter (1946) 3Scylaceus pallidus (Emerton, 1882) Cherryfi eld, Washington County litter, blueberry fi eld Maloney (2002)

Sisyrbe rustica (Banks, 1892) T18 MD BPP, Washington County litter, blueberry fi eld Drummond et al. (unpubl.)Walckenaeria breviaria (Crosby & Bishop, 1931) T18 MD BPP, Washington County litter, blueberry fi eld Drummond et al. (unpubl.)

 Walckenaeria castanea (Emerton, 1882) Steuben, Washington County litter, coastal red spruce stand HWRI (1991) 4

Winter Harbor, Hancock County litter, near fallen birch HWRI (2001) 4

Walckenaeria dondalei Millidge, 1983 T18 MD BPP, Washington County litter, blueberry fi eld Drummond et al. (unpubl.)

 Walckenaeria pinocchio (Kaston, 1945) Cherryfi eld & Columbia, Washington 

County

litter, blueberry fi eld Maloney (2002)

Walckenaeria tenella Millidge, 1983 Steuben, Washington County litter, coastal red spruce stand HWRI (1991) 4

Cherryfi ed, Washington County litter, blueberry fi eld Timms (unpubl.)

Walckenaeria sp. 10 T18 MD BPP, Washington County litter, blueberry fi eld Drummond et al. (unpubl.)

TETRAGNATHIDAE Pachygnatha tristriata C. L. Koch, 1845 Jonesboro, Washington County litter, blueberry fi eld Collins et al. (1996)

Columbia, Washington County litter, blueberry fi eld Drummond et al. (unpubl.)

Pachygnatha xanthostoma C. L. Koch, 1845 MDI, Hancock County (n. d.) 5 Procter (1946) 3

T18 MD BPP, Washington County litter, blueberry fi eld Drummond et al. (unpubl.)

Tetragnatha viridis Walckenaer, 1842 MDI, Hancock County (n. d.) 5 Procter (1946) 3

ARANEIDAE

Araneus corticarius (Emerton, 1884) MDI, Hancock County (n. d.) 5 Procter (1946) 3

Table 2.—continuedSpider taxa 1 Locality record(s) 2 Habitat(s) Source(s)

continued
182

Araneus thaddeus (Hentz, 1847) MDI, Hancock County (n. d.) 5 Procter (1946) 3Larinioides sclopetarius (Clerck, 1757) MDI, Hancock County (n. d.) 5 Procter (1946) 3

Steuben, Washington County lobster bait-house window HWRI (1991) 4

Mangora gibberosa (Hentz, 1847) MDI, Hancock County on patch of vegetation Procter (1946) 3

Winter Harbor, Hancock County search, hanging bog HWRI (1999) 4 Bar Harbor, Hancock County sweeping Kalmia-Vaccinium, 

grasses

Jennings & Cutler 

Bar Harbor, Hancock County sweeping vegetation, upland bog Jennings & Cutler 

(unpubl.)

Metepeira labyrinthea (Hentz, 1847) MDI, Hancock County heath Procter (1946) 3

Neoscona crucifera (Lucas, 1839) MDI, Hancock County (n. d.) 5 Procter (1946) 3Parazygiella montana (C. L. Koch, 1834) MDI, Hancock County branch of spruce tree Procter (1946) 3Zygiella x-notata (Clerck, 1757) MDI, Hancock County (n. d.) 5 Procter (1946) 3

DICTYNIDAE

Cicurina itasca Chamberlin & Ivie, 1940 T18 MD BPP, Washington County litter, blueberry fi eld Drummond et al. (unpubl.)Dictyna foliacea (Hentz, 1850) Columbia, Washington County litter, blueberry fi eld Maloney (2002)

T18 MD BPP, Washington County litter, blueberry fi eld Drummond et al. (unpubl.)

Emblyna annulipes (Blackwall, 1846) MDI, Hancock County (n. d.) 5 Procter (1946) 3Emblyna cruciata (Emerton, 1888) MDI, Hancock County (n. d.) 5 Procter (1946) 3

HUNTERS

DYSDERIDAE

Dysdera crocata C. L. Koch, 1838 MDI, Hancock County (n. d.) 5 Procter (1946) 3

MIMETIDAE

Ero canionis Chamberlin & Ivie, 1935 Cherryfi eld, Washington County litter, blueberry fi eld Maloney (2002)Ero leonina (Hentz, 1850) MDI, Hancock County sifting leaves Procter (1946) 3Mimetus epeiroides Emerton, 1882 MDI, Hancock County (n. d.) 5 Procter (1946) 3

LYCOSIDAE

Alopecosa kochi (Keyserling, 1877) Columbia & Jonesboro, Washington 

County

litter, blueberry fi eld Collins et al. (1996)

Cherryfi eld, Columbia, & T18 MD BPP, Washington County

litter, blueberry fi eld Drummond et al. (unpubl.)

Table 2.—continuedSpider taxa 1 Locality record(s) 2 Habitat(s) Source(s)

continued
183

Arctosa lama Dondale & Redner, 1983 Winter Harbor, Hancock County hanging bog HWRI (1999) 4

PMNWR, Steuben, Washington County

litter, maritime slope bog Jennings (unpubl.)

PMNWR, Steuben, Washington County

litter, Sphagnum bog Jennings (unpubl.)

Hogna helluo (Walckenaer, 1837) MDI, Hancock County (n. d.) 5 Procter (1946) 3

 Pardosa milvina (Hentz, 1844) Cherryfi eld, Washington County litter, blueberry fi eld Timms (unpubl.)

Columbia & T19 MD BPP, Washington County

litter, blueberry fi eld Collins et al. (1966)

Columbia, Washington County litter, blueberry fi eld Drummond et al. (unpubl.)

Pirata canadensis Dondale & Redner, 1981 Winter Harbor, Hancock County Sphagnum litter; hanging bog HWRI (1999) 4 Schizocosa avida (Walckenaer, 1837) T18 MD BPP & T19 MD BPP, 

Washington County

litter, blueberry fi eld Drummond et al. (unpubl.)

PISAURIDAE

Dolomedes scriptus Hentz, 1845 MDI, Hancock County female guarding nest, pond; dam Procter (1946) 3Dolomedes triton (Walckenaer, 1837) MDI, Hancock County from pond Procter (1946) 3Pisaurina mira (Walckenaer, 1837) Cherryfi eld, Washington County litter, blueberry fi eld Maloney (2002)

ANYPHAENIDAE

Wulfi la saltabundus (Hentz, 1847) T19 MD BPP, Washington County litter, blueberry fi eld Collins et al. (1996)

CLUBIONIDAE

Clubiona mixta Emerton, 1890 Cherryfi eld, Washington County litter, blueberry fi eld Maloney (2002)

Elaver excepta (L. Koch, 1866) MDI, Hancock County sifting (litter) Procter (1946) 3

CORINNIDAE

Phrurotimpus certus Gertsch, 1941 MDI, Hancock County (n. d.) 5 Procter (1946) 3

Amherst, Hancock County litter, blueberry fi eld Drummond et al. (unpubl.)Steuben, Washington County gravel pit, cobbles HWRI (1991) 4Steuben, Washington County litter, maritime slope bog Jennings (unpubl.)Jonesboro, Washington County litter, blueberry fi eld Maloney (2002)Cherryfi eld & Columbia, litter, blueberry fi eld Drummond et al. (unpubl.)

Phrurotimpus minutus (Banks, 1892) Cherryfi eld, Washington County litter, blueberry fi eld Drummond et al. (unpubl.)

Table 2.—continuedSpider taxa 1 Locality record(s) 2 Habitat(s) Source(s)

continued
184

GNAPHOSIDAE

Drassyllus depressus (Emerton, 1890) MDI, Hancock County sifting (litter) Procter (1946) 3

Columbia & Deblois, Washington County

litter, blueberry fi eld Collins et al. (1996)

T18 MD BPP, T19 MD BPP, Washington County

litter, blueberry fi eld Drummond et al. (unpubl.)

Drassyllus frigidus (Banks, 1892) Amherst, Hancock County litter, blueberry fi eld Drummond et al. (unpubl.)

Drassyllus rufulus (Banks, 1892) MDI, Hancock County (n. d.) 5 Procter (1946) 3

Gnaphosa brumalis Thorell, 1875 PMNWR, Steuben, Washington 

County

litter, maritime slope bog Jennings (unpubl.)

Micaria longispina Emerton, 1911 Columbia & T18 MD BPP, 

Washington County

litter, blueberry fi eld Drummond et al. (unpubl.)

Micaria riggsi Gertsch, 1942 Cherryfi eld, Columbia, Jonesboro, & 

T19 MD BPP, Washington County

litter, blueberry fi eld Collins et al. (1996)

Cherryfi eld, Columbia, & Jonesboro, Washington County

litter, blueberry fi eld Maloney (2002)

Columbia & T18 MD BPP, Washington County

litter, blueberry fi eld Drummond et al. (unpubl.)

Sergiolus decoratus Kaston, 1945 Cherryfi eld, Washington County litter, blueberry fi eld Collins et al. (1996)

Zelotes puritanus Chamberlin, 1922 Cherryfi eld, Columbia, & Jonesboro, 

Washington County

litter, blueberry fi eld Maloney (2002)

Cherryfi eld & Columbia, Washington County

litter, blueberry fi eld Drummond et al. (unpubl.)

PHILODROMIDAE

Ebo iviei Sauer & Platnick, 1972 Jonesboro, Washington County litter, blueberry fi eld Collins et al. (1996)

Jonesboro, Washington County litter, blueberry fi eld Maloney (2002)Cherryfi eld & T18 MD BPP, Washington County

litter, blueberry fi eld Drummond et al. (unpubl.)

Philodromus pernix Blackwall, 1846 MDI, Hancock County heath Procter (1946) 3

Cherryfi eld & Jonesboro, Washington County

litter, blueberry fi eld Maloney (2002)

T18 MD BPP, Washington County litter, blueberry fi eld Drummond et al. (unpubl.)

Philodromus histrio (Latreille, 1819) MDI, Hancock County (n. d.) 5 Procter (1946) 3

Table 2.—continuedSpider taxa 1 Locality record(s) 2 Habitat(s) Source(s)

continued
185

1 Within spider foraging guilds, enumeration of taxa follows Platnick (2007) “The world spider catalog, version 7.5.” New York: American Museum of Natural 

History, online at http://research.amnh.org/entomology/spiders/catalog/index.html

2 Locality source names as given in The Maine Atlas and GazetteerTM 29th ed. Yarmouth, ME: DeLorme. 78 p.3 Taxa listed by Procter (1946) have been updated to refl ect recent species synonymies and other taxonomic changes.4 HWRI, Humboldt Wildlife Research Institute, “Biology of Spiders” Seminar, Class of (year).5 (n. d.) = no data. 

Bar Harbor, MDI, Hancock County sweeping vegetation, upland bog Jennings & Cutler 

(unpubl.)

Columbia, Washington County litter, blueberry fi eld Drummond et al. (unpubl.)

Philodromus validus (Gertsch, 1933) Columbia & T18 MD BPP, 

Washington County

litter, blueberry fi eld Drummond et al. (unpubl.)

THOMISIDAE

Misumenoides formosipes (Walckenaer, 1837) MDI, Hancock County on fl owers, sweeping grass Procter (1946) 3

 Ozyptila americana Banks, 1895 MDI, Hancock County young from sifting leaves Procter (1946) 3

Xysticus discursans Keyserling, 1880 Cherryfi eld, Columbia, Jonesboro, & 

T19 MD BPP, Washington County

litter, blueberry fi eld Collins et al. (1996)

Cherryfi eld, Washington County litter, blueberry fi eld Timms (unpubl.)Cherryfi eld & Jonesboro, Washington County

litter, blueberry fi eld Maloney (2002)

Columbia, Washington County litter, blueberry fi eld Drummond et al. (unpubl.)

Xysticus gulosus Keyserling, 1880 MDI, Hancock County (n. d.) 5 Procter (1946) 3Xysticus fervidus Gertsch, 1953 T19 MD BPP, Washington County litter, blueberry fi eld Collins et al. (1996)

Cherryfi eld, Washington County litter, blueberry fi eld Maloney (2002)

Xysticus pellax O. Pickard-Cambridge, 1894 MDI, Hancock County (n. d.) 5 Procter (1946) 3

Deblois, Washington County litter, blueberry fi eld Collins et al. (1996)Jonesboro, Washington County litter, blueberry fi eld Maloney (2002)

SALTICIDAE

Habronattus waughi (Emerton, 1926) T18 MD BPP, Washington County litter, blueberry fi eld Drummond et al. (unpubl.)Naphrys pulex (Hentz, 1846) Jonesboro, Washington County litter, blueberry fi eld Maloney (2002)

Cherryfi eld, Washington County litter, blueberry fi eld Drummond et al. (unpubl.)

Pelegrina galathea (Walckenaer, 1837) MDI, Hancock County beating oak Procter (1946) 3

Tutelina elegans (Hentz, 1846) MDI, Hancock County heath Procter (1946) 3Tutelina formicaria (Emerton, 1891) MDI, Hancock County (n. d.) 5 Procter (1946) 3

Table 2.—continuedSpider taxa 1 Locality record(s) 2 Habitat(s) Source(s)
186

Loss of habitat and invasion by exotic species are the two leading causes of species endangerment 

(Czech et al. 2000, Wilson 1992). Unfortunately, like most invertebrates, spiders are not immune to 

such adverse effects on species richness and diversity. For example, drainage of freshwater marshes at 

Milbridge could adversely impact resident spider species such as Antistea brunnea, Clubiona norvegica, 

and Philodromus peninsulans. Spiders associated with seashore cobble beaches (e.g., Diplostyla 

concolor, Pardosa groenlandica, Habronattus borealis) are susceptible and vulnerable to maritime oil 

spills. Apparently, some native spiders at Milbridge are already being displaced by invasive aliens such 

as Achaearanea tabulata and Steatoda bipunctata; most likely others will be impacted by Linyphia 

triangularis, an aggressive species recently introduced from the Palearctic (Jennings et al. 2002). 

Because of these and other potential threats, and the importance of spiders as bioregulators (Nyffeler 

and Benz 1987, Nyffeler et al. 1994, Riechert and Lockley 1984), and as bio-indicators of habitat 

quality (Allred 1975, Clausen 1986, Marc et al. 1999, Vollrath 1988), spiders warrant much greater 

attention from wildlife, land, and conservation managers. Perhaps this study will provide a basis for 

future evaluations and assessments of the araneofauna, and its health, at Milbridge.

ACKNOWLEDGMENTS

Portions of this research were done in collaboration with the U. S. Department of the Interior, Fish 

and Wildlife Service, Maine Coastal Islands National Wildlife Refuge, formerly the Petit Manan 

National Wildlife Refuge. We greatly appreciate the cooperative assistance of refuge biologists Linda 

J. Welch and Michael C. Langlois for their logistical support, training and supervision of summer 

interns, and other invaluable contributions. Refuge student interns Michelle Fisher, Barry Southard, 

and Bret Irving assisted with collection of fi eld samples and initial sortings of specimens; we thank 

them for their dedicated service.

Students of the “Biology of Spiders” seminars taught by Dr. Kefyn M. Catley at the Humboldt 

Wildlife Research Institute, Steuben, ME, shared specimens and collection data from Milbridge. Dr. 

Francis A. Drummond and associates, University of Maine, Orono, provided numerous unpublished 

records of spiders associated with commercial blueberry fi elds in Maine’s “downeast” region; we 

gratefully acknowledge their cooperative assistance.

Dr. Bruce Cutler, University of Kansas, Lawrence, assisted with collections on Bois Bubert Island 

in 2001. His expertise and superior collecting skills yielded several new State records of species 

previously unknown in Maine. We thank Arlene Banks, James Bird, and Mel Johnson, Fogler Library, 

University of Maine, Orono, for their expert literature search and retrieval services.

Michael C. Langlois prepared Figure 1; Cynthia A. Jennings and Diane L. Rowland provided much 

needed word-processing assistance; and Nancy B. Jennings assisted with data summaries and literature 

citations. We are grateful for their help.

We give special thanks to our manuscript reviewers for their constructive comments and suggestions: 

Drs. Paula E. Cushing, Denver Museum of Nature and Science, Denver, CO; Charles D. Dondale, 

Biodiversity Research Branch, Agriculture and Agri-Food Canada, Ottawa, ON; and K. Elizabeth 

Gibbs, Department of Biological Sciences, University of Maine, Orono. We greatly appreciate their 

devoted time and efforts.
187

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Common Name Latin Name

Alder, speckled Alnus incana (L.) Moench ssp. rugosa (Du Roi) Clausen

Apple Malus sp.

Aspen, bigtooth Populus grandidentata Michx.

Aspen, quaking (trembling) Populus tremuloides Michx.

Aster Aster sp.

Astilbe Astilbe sp.

Balsam fi r Abies balsamea (L.) Mill.

Bayberry Myrica pensylvanica Loisel.

Birch Betula sp.

Birch, gray Betula populifolia Marsh.

Birch, white (paper) Betula papyrifera Marsh.

Blackberry Rubus sp.

Blueberry Vaccinium sp.

Blueberry, common lowbush Vaccinium angustifolium Ait.

Blue-joint grass Calamagrostis canadensis (Michx.) Beau.

Bush-honeysuckle Diervilla lonicera P. Mill.

Cedar, Northern white Thuja occidentalis L.

Cherry, pin Prunus pensylvanica L. f.

Cherry, wild Prunus sp.

Chokecherry Prunus virginiana L.

Cinquefoil Potentilla sp.

Clover, white and yellow Melilotus offi cinalis (L.) Lam.

Crabapple Malus prunifolia (Willd.) Borkh.

Crowberry, black Empetrum nigrum L.

Daisy, ornamental Leucanthemum sp.

Daisy, ox-eye Leucanthemum vulgare Lam.

Dunegrass, American Leymus mollis (Trin.) Hara

Garden-heliotrope Valeriana offi cinalis L.

Goldenrod Solidago sp.

Goldenrod, common Solidago canadensis L.

Goldenrod, seaside Solidago sempervirens L.

Hawkweed, meadow Hieracium caespitosum Dumort.

Hawthorn Crataegus sp.

APPENDIX

Common and Latin names of Plants with Associated Spiders, 

Milbridge, Washington County, Maine. 1

continued
204

Larch, eastern Larix laricina (Du Roi) K. Koch

Leatherleaf Chamaedaphne calyculata (L.) Moench

Lilac Syringa sp.

Maple, red Acer rubrum L.

Meadowsweet Spiraea alba (Du Roi) var. latifolia (Ait.) Dippel

Mountain-ash Sorbus americana Marsh.

Oak, Northern red Quercus rubra L.

Pea, common Pisum sativum L.

Potato Solanum tuberosum L.

Pine, jack Pinus banksiana Lamb.

Pine, white Pinus strobus L.

Raspberry Rubus idaeus L.

Reindeer lichen Cladina rangiferina (L.) Nyl.

Rhodora Rhododendron canadense (L.) Torr.

Rockweed Fucus vesiculosus L.

Rose, rugosa Rosa rugosa Thomb.

Rush, black Juncus gerardii Loisel.

Russian olive Elaeagnus angustifolia L.

Sarsaparilla, wild Aralia nudicaulis L.

Seaside-pea Lathyrus japonicus Willd.

Sedge Scirpus sp.

Serviceberry Amelanchier sp.

Sheep laurel Kalmia angustifolia L.

Spruce, red Picea rubens Sarg.

Spruce, white Picea glauca (Moench) Voss

St. Johnswort, common Hypericum perforatum L.

Sunfl ower Helianthus sp.

Sweet gale Myrica gale L.

Sweetpea Lathyrus sp.

Timothy Phleum pratense L.

Wild-raisin Viburnum nudum var. cassinoides (L.) Torr. & Gray

Willow Salix sp.

Winterberry Ilex verticillata (L.) Gray

Witch alder Fothergilla gardenii L.

1 Most of the common and Latin names are based on: Haines, A.; Vining, T.F. 1998. Flora of 

Maine: a manual for identifi cation of native and naturalized vascular plants of Maine. V.F. Thomas 

Company: Bar Harbor, ME: 847 p.

Appendix—continued
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Jennings, Daniel T.; Graham, Frank Jr. 2007. Spiders (Arachnida: Araneae) Of 

Milbridge, Washington County, Maine. Gen. Tech. Rep. NRS-16. Newtown 

Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research 

Station. 204 p.

An inventory or spiders associated with diverse habitats of Milbridge, a 6,290-ha 

area of the East Coastal BioPhysical Region, yielded 6,979 individuals of 19 families, 

145 genera, and 302 species (4 unknown). Species richness per genus ranged from 

1 to 13, with 88 genera represented by a single species. Total species composition 

favored web spinners over hunters; however, more hunters were collected than 

web spinners, and more females were collected than males. Pitfall traps yielded the 

greatest number of species and individuals; fully 47 percent of the inventoried spiders 

were method-unique species. Species-faunal compositions were generally distinct 

among the sampled habitats and included habitat-unique species. Species-range 

extensions include new state, regional, and national records. The total estimated 

araneofaunal richness for Milbridge is 24 families, 172 genera, and ~ 411 species.

KEY WORDS: spider faunal inventories, species richness, sampling methods, 

spider-habitat associations, method-unique species, habitat-unique species, invasive 

species, estimated faunal richness
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