3 S TEXAS TECH UNIVERSITY Natural Science Research Laboratory Special Publications Museum of Texas Tech University Number 67 17 July 2018 Harvestmen of the Family Phalangiidae (Arachnida, Opiliones) in the Americas James C. Cokendolpher and Robert G. Holmberg Front cover: Opilio parietinus in copula (male on left with thicker legs and more spines) from Baptiste Lake, Athabasca County, Alberta. Photograph by Robert G. Holmberg. Special Publications Museum of Texas Tech University Number 67 Harvestmen of the Family Phalangiidae (Arachnida, Opiliones) in the Americas James C. Cokendolpher and Robert G. Holmberg Layout and Design: Cover Design: Production Editor: Lisa Bradley Photograph by Robert G. Holmberg Lisa Bradley Copyright 2018, Museum of Texas Tech University This publication is available free of charge in PDF format from the website of the Natural Sciences Research Laboratory, Museum of Texas Tech University (nsrl.ttu.edu). The authors and the Museum of Texas Tech University hereby grant permission to interested parties to download or print this publication for personal or educational (not for profit) use. Re-publication of any part of this paper in other works is not permitted without prior written permission of the Museum of Texas Tech University. This book was set in Times New Roman and printed on acid-free paper that meets the guidelines for per¬ manence and durability of the Committee on Production Guidelines for Book Longevity of the Council on Library Resources. Printed: 17 July 2018 Library of Congress Cataloging-in-Publication Data Special Publications of the Museum of Texas Tech University, Number 67 Series Editor: Robert D. Bradley Harvestmen of the Family Phalangiidae (Arachnida, Opiliones) in the Americas James C. Cokendolpher and Robert G. Holmberg ISSN 0149-1768 ISBN 1-929330-34-0 ISBN13 978-1-929330-34-8 Museum of Texas Tech University Lubbock, TX 79409-3191 USA (806)742-2442 Harvestmen of the Family Phalangiidae (Arachnida, Opiliones) in the Americas James C. Cokendolpher and Robert G. Holmberg Abstract This paper reviews 68 scientific names of species of Phalangiidae reported from North, Central, and South America. We found that: 1)16 species of Phalangiidae distributed across 10 genera ( Leptobunus , Liopilio , Lophopilio , Mitopus, Odiellus , Oligolophus, Opilio, Paroligolophus, Phalangium, and Rilaena ) are still valid; 2) up to 38% of these species have been introduced into North America from Europe and pos¬ sibly Asia; and 3) no members of this family are native to Mexico, Central and South America, or the Caribbean. An additional genus and species (i.e., Megabunus diadema) is recorded from Iceland and is to be watched for in North America. We recognize four new synonyms, one change in family status, and two other changes in status. We confirm both the statement by Levi and Levi (1955) that Opilio angulatichelis Roewer 1952 = Phalangium opilio Linnaeus 1758 and Bishop’s (1949) conclusion that Lacinius ohioensis (Weed 1889a) = Odielluspictus (Wood 1868). Egaenus and Lacinius do not occur anywhere in the Americas. Range extensions (i.e., new provinces or states) are given for Liopilio glaber (1 new extension), Lophopilio palpinalis (1, introduced; new record for North America), Mitopus morio (2 new provinces), Odiellus pictus (3 new provinces; 1 new USA county), Rilaena triangularis (1, introduced), Opilioparietinus (2, possibly introduced), and Phalangium opilio (18, possibly introduced). Taxonomic changes are: lectotypes and paralectotypes are designated for Mitopus californicus Banks 1895, Odiellus pictus argentus Edgar 1966, and Oligolophus montanus Banks 1893c; Lacinius texanus Banks 1893a = Odiellus pictus (Wood 1868), new synonym; Mitopus californicus Banks 1895 = Opilio parietinus (De Geer 1778), new synonym; Mitopus dorsalis Banks 1900 = Mitopus morio Pabricius (1779), new synonym; Paro- diellus projectus Goodnight and Goodnight 1942a (Phalangiidae) = Protolophus niger Goodnight and Goodnight 1942b (Protolophidae), new synonym, new family rank; Opilio chickeringi = Lanthanopilio chickeringi is placed in the “Metopilio group”, new status; and Phalangium rudipalpe Gervais 1849 = Genus and species inquirenda (Sclerosomatidae: Gagrellinae), new status. Anew subgeneric name, Slimlepto , is given for the parvulus group of Leptobunus. A taxonomic key to the valid Phalangiidae of the Americas is provided. Key words: Leptobunus , Liopilio , Lophopilio , Megabunus , Mitopus , Odiellus , Oligolophus , Opilio , Paroligolophus , Phalangium , Rilaena , Slimlepto Introduction The Opiliones (formerly, also referred to as the Phalangida) are the third largest order of the class Arachnida. Colloquially they are known in the Americas as the daddy-long-legs or the more acceptable name of harvestmen. With slightly over 6,650 species worldwide, the Opiliones are split unequally among four suborders (Kury 2017). Almost all long-legged harvestmen observed walking or resting in/on open situations like leaves, bark, walls, etc. (both during the daytime and night) will be members of the suborder Eupnoi. The most common and conspicuous members of this suborder will be the species of the Phalangiidae and Sclerosomatidae. 1 2 Special Publications, Museum of Texas Tech University The Phalangiidae (with over 380 species) occur throughout most of the cold temperate parts of the Northern Hemisphere. In these areas, they are the most likely family of harvestmen encountered by humans in disturbed areas such as croplands, gardens, and urban areas. However, some species, and most other Opil- iones families, are found only in undisturbed habitats of forests and mountains. Phalangiids are moderately sized, with globular bodies and thin legs, not heavily sclerotized, and drab colored. For this study, we include all the Nearctic and Neotropical Realms of the Americas, including the Caribbean islands, as well as Kalaallit Nunaat (Green¬ land) and Iceland. There are three or four Phalangiidae species on Iceland. Henriksen (1932), Stol (2007), and Bezdecka et al. (2017) recorded Megabunus diadema (Fabricius 1779), Mitopus morio (Fabricius 1779), and Oligolophus tridens (C. L. Koch 1836) (as well as Nemastoma bimaculatum (Fabricius 1775, Family Ne- mastomatidae). However, Spoek (1963) also mentions Opilio parietinus (De Geer 1778) as being present in Iceland but gave no further information. All these spe¬ cies occur further to the west in North America except Megabunus. Iceland is part of the mid-Atlantic ridge which is the boundary between the North American and Eurasian plates. As it originated volcanically from the collision of the two plates, is far from other land masses, and phalangiids have poor dispersal capabili¬ ties, the Opiliones of Iceland probably resulted from human activities and we predict that Megabunus will eventually populate more of North America. Thus, it is included in this paper. Shear (2016) reviewed the known Eurasian Opiliones species that have been recorded as being introduced into North America: Nemastoma bi¬ maculatum (Fabricius 1775) (Nemastomatidae from Europe) and Trogulus tricarinatus (Linnaeus 1767) (Trogulidae from Europe). He also recorded three European members of the Phalangiidae that have been introduced: Oligolophus tridens (C. L. Koch 1836), Paroligolophus agrestis (Meade 1855), and Rilaena triangularis (Herbst 1799). He also briefly mentioned that three other species might have been introduced but might otherwise be of a natural Holarctic distribution: Mitopus morio (Fabricius 1779, Opilio parietinus (De Geer 1778), and Phalangium opilio (Linnaeus 1758). It is clear that the former three phalangiid species are recent introductions to the New World from Europe. In this publication, we also add another species of Pha¬ langiidae introduced from Europe into North America: Lophopilio palpinalis (Herbst 1799). The purpose of this study was to document the identity and distributions of, and provide a means to identify, members of the Phalangiidae in the Americas. Where possible we have included data on the juvenile life stage as well as the adults. Phalangium Linnaeus 1758 (originally Greek) and Opilio Herbst 1798 (origi¬ nally Latin) have appeared in the taxonomic literature many times and in combinations with what are now understood not to be closely related species (some even in separate families). All of these combinations based on species reported from the Americas are reviewed and updated where needed. The member species of the genera Odiellus Roewer 1923 and Mitopus Thorell 1876 are distributed throughout the Holarctic region and both are in need of modem taxonomic studies. The last times they were reviewed on a world basis was by Roewer 1956 and 1957. At that time, the morphology of the genitalia and genetics were not studied. Systematics ORDER OPILIONES Sundevall 1833 Suborder Eupnoi Hansen and Sorensen 1904 Superfamily Phalangioidea Latreille 1802 Family Phalangiidae Latreille 1802 Because Phalangium was the first genus of Opil¬ iones to be named by Linnaeus in 1758, its definitions have changed remarkably over the centuries. Likewise, the family based upon that genus has shown great taxonomic changes over time. For this reason there are numerous species that were named early-on in combination with Phalangium that are no longer even recognized in the same families or, occasionally, the same suborders. The current use of the Phalangiidae COKENDOLPHER AND HOLMBERG-PHALANGIIDAE OF THE AMERICAS 3 has been fairly constant since about the middle 1970’s, but there is some disagreement regarding the number and position of the subfamilies (Hedin et al. 2012; JCC personal observation). Kury (2013) estimated 55 extant genera with 394 extant species in the Phalangiidae. The family is characterized by a combination of characters recorded below, but only that of the male genitalia is uniquely diagnostic: palp, leg-like but may be armed ventrally with spines or spine-tipped tubercles; palpal tarsus longer than the palpal tibia; palpal claw single, well developed, curved, simple (smooth, without bumps, denticulation/pectination); palps and chelicerae clearly visible from above, not covered by a hood; eyes raised above the prosoma on an ocularium, spination vari¬ able; leg coxae without denticles (especially on lateral margins); single, undivided claw on terminus of each leg; second leg (leg II) usually longest; no nodules or pseudosegments in any leg femora; genitalia covered with a ventral operculum that extends backwards from the posterior edge of the second leg coxae to beyond the fourth leg coxae, operculum flexible, not heavily sclerotized; penis, in lateral view, strongly bent between the long corpus and shorter glans; stylus simple and not twisted; ovipositor long, many segmented, and flexible. It is not clear what characters best separate the subfamilies (Martens 1978; Crawford 1992). For the 11 genera considered valid for the Americas, we placed them according to Crawford 1992 and will discuss them in this order: Phalangiinae Latreille 1802: Leptobunus, Liopilio, Phalangium Opilioninae C. L. Koch 1839a: Opilio Oligolophinae Banks 1893b: Mitopus, Odiellus, Oligolophus, Paroligolophus Platybuninae Star^ga 1976: Lophopilio, Mega- bunus, Rilaena Misplaced species: Parodiellus Subfamily Phalangiinae Latreille 1802 Leptobunus Banks 1893b (Figs. 1-13) Cokendolpher (1985) revised the taxonomy of this genus. He recognized two parts: the californicus group and the parvulus group. We propose here to recognize and formalize the naming of the californicus group in the nominal subgenus Leptobunus ( Leptobu¬ nus ) because the type species of the genus is included in this group. The second group we propose to call Leptobunus ( Slimlepto). Slimlepto, new subgenus, is fitting as the first part of the new name is “slim”, in English to be slender, and the second part, “lepto” is a shortened form of the genus Leptobunus. In this case, we are not using “lepto” as a Greek adjective meaning thin. Slimlepto is considered masculine and therefore the endings of the existing species names remain the same: Leptobunus ( Slimlepto)parvulus , type species, new combination, and Leptobunus {Slimlepto) aureus , new combination. The subgenera can be distinguished by the differences noted in the key and illustrations. Leptobunus aureus Cokendolpher 1985 was reported only from California. Leptobunus borealis Banks 1898a has been collected from Bering and Copper Islands of Russia as well as coastal Alaska. Leptobunus californicus Banks 1893b was reported from California and Nevada. Schenkel (1951) named Liomitopus laevis; however, Cokendolpher (1980a) synonymized it under L. californicus. Leptobunus parvulus (Banks 1894) (= Liobunumparvulum, Leuro- ny chus parvulus) is distributed from Alaska to Califor¬ nia (Cokendolpher 1985; Bragg and Holmberg 2009). A fossil harvestman described from the USA (Colorado) as Leptobunus atavus Cockerell 1907 has been transferred to the genus Amauropilio Mello- Leitao 1937 by Cokendolpher and Cokendolpher 1982. Similarly, the Mexican (Michoacan) species described as Leptobunus mexicanus Goodnight and Goodnight 1942c was moved to the genus Paranelima Caporiacco 1938 by Cokendolpher (1985). Both of these species are now placed in the Sclerosomatidae: Leiobuninae (Cokendolpher and Lee 1993). Leptobunus spinula- tus Banks 1898b, described from Mexico (Nayarit, Guanajuato, Jalisco, Mexico, and Michoacan), has 4 Special Publications, Museum of Texas Tech University Figure 1. Leptobunus californicus dorsal view of male palpal tibia and tarsus that are noticeably expanded on mesal margins, setae not illustrated. Illustration from Cokendolpher 1985. Figure 2. Leptobunus californicus male palp, mesal view. Illustration from Cokendolpher 1985; flipped horizontally so palp faces left. Figure 3. Leptobunus parvulus mesal view of male chelicera. Illustration from Cokendolpher 1985. Figure 5. Leptobunus parvulus lateral view of distal end of penis. Illustration from Cokendolpher 1985. Figure 6. Leptobunus parvulus lateral view of seminal receptacle. Illustration from Cokendolpher 1985. Figure 7. Leptobunus californicus lateral view of distal end of penis. Illustration from Cokendolpher 1985. Figure 4. Leptobunus aureus lateral view of distal end of penis. Illustration from Cokendolpher 1985. COKENDOLPHER AND HOLMBERG-PHALANGIIDAE OF THE AMERICAS 5 Figure 8. Leptobunus californicus lateral view of seminal receptacle. Illustration from Cokendolpher 1985. Figure 9. Leptobunus borealis lateral view of seminal receptacle. Illustration from Cokendolpher 1985. Figure 11. Leptobunus borealis dorsal view of male palpal tibia and tarsus; setae not illustrated. Illustration from Cokendolpher 1985. Figure 12. Leptobunus borealis lateral view of distal end of penis. Illustration from Cokendolpher 1985. Figure 13. Preserved Leptobunus borealis , sex not determined, from Kenai National Wildlife Refuge, Alaska. Photograph by Matt Bowser. Figure 10. Preserved Leptobunusparvulus female from Vancouver Island, British Columbia. Photograph by Don J. Buckle. 6 Special Publications, Museum of Texas Tech University been transferred to Metopilio Roewer 1911 by Coken- dolpher (1985). This genus currently is placed in an undescribed assemblage (new family) that is informally referred to as the “ Metopilio group” (Cokendolpher and Lee 1993; Hedin et al. 2012). This group provision¬ ally has been treated in the Sclerosomatidae (Tourinho 2007), but because of the striking differences in penial morphology (JCC personal observations) and molecu¬ lar studies of Opiliones (Giribet et al. 2002; Hedin et al. 2012), it appears to be distinct from both Scleroso¬ matidae and Phalangiidae. Liopilio Schenkel 1951 (Figs. 14-21) Cokendolpher and Lee (1993) reported that Lio¬ pilio glaber Schenkel 1951 occurred in the mountains of Alberta, British Columbia, and Washington. Anew state record is: USA: MONTANA: Park County: Grass¬ hopper Glacier, 24 km North of Cooke City, 21 August 1959, D. C. Lowerie, 1 female (AMNH); Stillwater County: “W. Rosebud Plat.” [Plateau?], 21 July 1966, B. and C. Durden, 1 female (AMNH). Both of these records are from 3,050-3,200 m elevation. Cokendolpher (1981a) described Liopilio yukon from Yukon Territory and Alaska. We have no new distribution records of this species. Figure 14. Liopilio yukon ventral view of distal end of male palp; setae not illustrated. Illustration from Cokendolpher 1981a. Figure 15. Liopilio yukon dorsal view of distal end of female palp. Illustration from Cokendolpher 1981a. Figure 16. Liopilio yukon dorsal view of male body. Illustration from Cokendolpher 1981a. Figure 17. Liopilio glaber dorsal view of male body. Illustration from Cokendolpher 1981a. Figure 18. Live Liopilio glaber male from Lemming Lake, Kakwa Wildlife Park, Alberta. Photograph by Robert G. Holmberg; enlargement of Fig. 20. COKENDOLPHER AND HOLMBERG-PHALANGIIDAE OF THE AMERICAS 7 Figure 19. Live Liopilio glaber female from Lemming Lake, Kakwa Wildlife Park, Alberta. Photograph by Robert G. Holmberg; rotated enlargement of Fig. 21. Figure 20. Live Liopilio glaber male from Lemming Lake, Kakwa Wildlife Park, Alberta. Photograph by Robert G. Holmberg. Figure 21. Live Liopilio glaber female from Lemming Lake, Kakwa Wildlife Park, Alberta. Photograph by Robert G. Holmberg. Phalangium Linnaeus 1758 (Figs. 22-26) Because of the antiquity of the name, some of the older specific names to New World harvestmen have been associated incorrectly with Phalangium and the Phalangiidae, namely those of Say (1821), Gervais (1849), and Wood (1868, 1869). The named Phalangium from the New World are arranged alpha¬ betically below. Only a few references are recorded for each species (the original and usually one or two more relevant ones). Phalangium acanthipes Pickard-Cambridge 1905, from Guerrero, Mexico, in Metopilio by Roewer (1911) (new family, “ Metopilio group”). Phalangium armigerum Pickard-Cambridge 1905 from Guerrero and Morelos, Mexico, in Metopilio by Roewer (1911) (new family, “ Metopilio group”). Phalangium bicolor Wood 1868 (^ Phalangium bicolor Fabricius 1793; Wood’s name is a primary homonym of that named by Fabricius and as such it was replaced by the oldest available synonym, Leiobunum elegans Weed 1889b; see Cokendolpher 1984) = Ne- lima elegans (Weed 1889b); Cokendolpher and Zeiders (2004) in Sclerosomatidae: Leiobuninae. Phalangium calcar = Leiobunum calcar (Wood 1868), Davis (1934), Ingianni et al. (2011) in Sclero¬ somatidae. Phalangium cinereum Wood 1868 = Opilio parietinus (De Geer 1778)—see below under Opilio. Phalangium dorsatum = Leiobunum vittatum (Say 1821); Davis (1934) in Sclerosomatidae. Phalangium exilipes = Leiobunum exilipes (Wood 1868); Davis (1934) in Sclerosomatidae. Hallan (2005) recorded in his internet biology catalog the species name spelled with two l’s and incorrectly recorded the year of description as 1878. Unfortunately, this has been followed generally by internet related catalogs and other resources. It is incorrect and a misspelling. Wood (1868; page 23) clearly spelled the name with a single 1. 8 Special Publications, Museum of Texas Tech University Figure 22. Phalangium opilio prosoma including ocularium and eyes, ventral arrow points to left ozopore, center arrow points to spine-tipped spur on left supracheliceral laminae, sex not determined. SEM photograph by Robert G. Holmberg. Figure 23. Phalangium opilio lateral view of male. Illustration traced and modified from Bishop 1949; flipped horizontally so body faces left. Figure 24. Phalangium opilio dorsal body view, chelicera, and palp of male. Illustration traced from Bishop 1949. Figure 25. Live Phalangium opilio male from Saskatoon, Saskatchewan. Photograph by Don J Buckle. Figure 26. Live Phalangium opilio female from Ottawa, Ontario. Photograph by Robert G. Holmberg, modified; flipped horizontally so body faces left. COKENDOLPHER AND HOLMBERG-PHALANGIIDAE OF THE AMERICAS 9 Phalangium favosum Wood 1868 = Trachyrhi- nus favosus (Wood 1868); Cokendolpher (1981b) in Sclerosomatidae. Phalangium formosum = Leiobunum formosum (Wood 1868); Davis (1934) in Sclerosomatidae. Shultz (2012) noted that this species was “soon to be trans¬ ferred to Hadrobunus .” Phalangium grande = Hadrobununs grandis (Say 1821); Shultz (2010, 2012) in Sclerosomatidae. Phalangium horridum Pickard-Cambridge 1905 from Guerrero and Michoacan, Mexico, is a secondary homonym of the Old World species P. horridum Panzer 1794 (now used in the combination Lacinius horridus, Phalangiidae). Pickard-Cambridge’s P horridum was placed in Metopilio by Roewer (1911) (new family, “Metopilio group”). Because it is a homonym, Meto¬ pilio cambridgei Mello-Leitao 1944 was supplied as the replacement name. Phalangium longipalpis — see below under Pha¬ langium opilio. Phalangium maculatipes Pickard-Cambridge 1905, from Guerrero, Hidalgo, and Puebla, Mexico, was placed in Metopilio by Roewer (1911) (new family, “Metopilio group”). Phalangium maculosum = Hadrobunus maculo- sus (Wood 1868); Shultz (2010) in Sclerosomatidae. Phalangium nigropalpi = Leiobunum nigropalpi (Wood 1868); Davis (1934), Ingianni (2010) in Sclero¬ somatidae. Phalangium nigrum = Eumesosoma nigrum (Say 1821); Cokendolpher (1980b) in Sclerosomatidae. Phalangium opilio Linnaeus 1758 was the first harvestman officially named, with types from Europe and, later (as synonyms), North America (Crawford 1992; Cokendolpher and Lee 1993). It is the only member of the genus currently recorded from North America. Phalangium opilio is found throughout much of the Holarctic region as well as New Zealand (Martens 1978; Butcher etal. 1988) and Japan (Suzuki and Tsurusaki 1983; Tsurusaki and Takenaka 2010). Phalangium opilio is now the most widely distributed species of harvestmen in the world (Novak et al. 2009). It was first recorded in North America as Phalangium longipalpus Weed 1890 and later as P. opilio and Opilio angulatichelis Roewer 1952 (see below under Opilio ). As human activities are suspected for its transport to Newfoundland (Lindroth 1957; p. 190), New Zealand (Gruber and Hunt 1973), and Japan (Tsurusaki and Tak¬ enaka 2010), and this species usually is found in areas disturbed by humans but very seldom in undisturbed habitats, we support the conclusion that this com¬ mon and very wide spread species is not native to the Americas (Bragg and Holmberg 2009). Cokendolpher and Lee (1993) recorded P. opilio from five provinces of Canada (Alberta, British Columbia, both New¬ foundland and Labrador, Ontario, and Saskatchewan) and 17 states of the USA (Alaska, Arkansas, Idaho, Illinois, Maine, Massachusetts, Michigan, Minnesota, Montana, New York, North Dakota, Ohio, Oklahoma, Pennsylvania, Texas, Washington, and Wisconsin). Koponen (1995) recorded it from Quebec, Newton and Yeargan (2002) from Kentucky, and Clark et al. (1994) from Virginia. Six new provincial/territorial and 13 new state/ district records are: CANADA: MANITOBA: Birch River campground, Primrose Wayside Park, 25 July 1997, RGH, 1 male (RGH); Blue Lake campground, Duck Mountain Provincial Park, 15 August 1969, ROM field Party, 1 male (ROM); 4 km west of Clear Lake, Riding Mountain National Park, 21-30 August 1979, R. B. Sample, 3 males, 9 females (CNC); Delta Marsh, University of Manitoba field Station, summer 1970, L. Hlynka, 2 males (RGH); 18 km north on Highway 10 of Lorrest, 5 September 1970, ROM field Party, 1 male (ROM); Glen Lea Research Station, 16 km south of Winnipeg, 8-15 September 1972, R. G. Sample, many males and females (CNC); Grandview Beach, 11-12 August 1970, ROM field Party, 2 males, 1 fe¬ male (ROM); Lake Audy, Riding Mountain National Park, 21-28 August 1979, J. and M. Redner, 3 males, 5 females (CNC); Lake of the Prairies, 24 July 1997, RGH, 1 male (RGH); 59 km north of Mafeking, 16 August 1969, ROM field Party, 1 female (ROM); 20 km south of Melita, 23 July 1997, RGH, 1 female (RGH); Moon Lake, Riding Mountain National Park, 9 September 1987, CWA, 1 female (CWA); Swanson 10 Special Publications, Museum of Texas Tech University Spring, Riding Mountain National Park, several col¬ lections 29-30 August 1979, D. B. Lyons and S. J. Miller, 8 males, 24 females (CNC); 15 km southwest of Swan River, 25 July 1997, RGH, 1 male (RGH); Virden and 40 km southwest, 23-24 July 1997, RGH, 2 males, 2 females (RGH). NEW BRUNSWICK: Fredericton, 11-16 July 1978, C. D. Dondale, 1 male (CNC); Kouchibouquac National Park, several collec¬ tions 12 July-12 September 1977, several collectors, many males and females (CNC); St Andrews, 17 July 1978, S. A. Marshall, 1 male (CNC); Saint John, 20 July 1968, E. E. Linquist, 2 males (CNC); St-Leonard, 23 September 1974, C. D. Dondale, 1 female (CNC). NORTHWEST TERRITORIES: campsite at NWT- Alberta border (60°00’N, 116°59 , W), 13-14 August 1998, RGH, 1 male (RGH); Fort Liard, summer 2000, R. Smith, 3 males, 4 females (RGH); Fort Smith, 16 August 1998, RGH, 1 male, 1 female (RGH); Hay River, 15-19 August 1998, RGH, 1 male (RGH); Yellowknife, 18-30 July 2009, summer 2010, and 15 July-6 August 2011, Brian Latham, 28 males, 8 females (RGH). NOVA SCOTIA: Barton, 23 July 1959, C. D. Dondale, 1 male, 1 female (CNC); Brair Island, 20 September 1996, DJB, 3 males (DJB); Canard, vari¬ ous dates between 1956 and 1959, C. D. Dondale, 8 males, 7 females (CNC); Coldbrook, 10-11 July 1957 and 1959, C. D. Dondale, 2 females (CNC); Cow Bay, 28 September 1956, C. D. Dondale, 1 male, 1 female (CNC); Digby, 29 July-23 August 1960, Fox and Sawler, many males and females (CNC); Graywood, 1-12 July 1957, unknown collector, many males and females (CNC); Gulliver’s Cove, 20 September 1996, DJB,1 male (DJB); Halls Harbour, 27 July-31 August 1960, unknown collector, many males and females (CNC); Kentville, 29 August and 3 September 1966, C. D. Dondale, 4 males, 3 females (CNC); Lakeview, 4 October 1956, unknown collector, 5 males, 1 female (CNC); Lily Lake, 27 September 1960, C. D. Dondale, 1 female (CNC); Marshalltown, 14 September-5 Octo¬ ber 1960, unknown collector, many males and females (CNC); Martins River, 27 July 1956, C. D. Dondale, 1 male (CNC); Sable Island, several collections between 13 July-15 September 1967, J. E. H. Martin, several males and females (CNC); Sable Island, 8-12 Septem¬ ber 1977, B. Wright, 1 male (CNC); Sweets Corner, 14 August 1956, C. D. Dondale, 1 female (CNC); White Rock, 14 September 1956, C. D. Dondale, 8 males (CNC). PRINCE EDWARD ISLAND: Malpeque Bay, Green Park, 5-8 September 1972, C. Starr, 1 male, 1 female (CNC). YUKON: Carcross, 14 August 1987, RGH, 3 males, 1 female (RGH); Takhini Hot Springs, 20 km northwest of Whitehorse, 16 August 1987, RGH, 3 males, 3 females (RGH); 5 km northwest of Watson Lake, 17 August 1987, RGH, 1 male (RGH); White¬ horse, 25 August 1969, J. Belicek, 1 male, 1 female (RGH); Whitehorse, 9 August 1982, J. J. Robinson, 1 male (UBC); Whitehorse, 10 August 1987, RGH, 3 males, 1 female (RGH). USA: ALASKA: Municipal¬ ity of Anchorage: Anchorage, 100 m, August 1974, N. L. H. Krauss, 2 females (AMNH). Yukon-Koyukuk: Circle Hot Springs, 65.29°N, 144.40°W, 14 August 1968, W. Ivie, 1 female (AMNH). CALIFORNIA: Contra Costa County: Richmond, 15 January-15 March 1987, S. Stockwell, J. Steele, 2 males, 1 female (JCC); Monterey County, Monterey, 5 June 1982, R. Johnson, Jr., 4 males, 4 females (JCC); Riverside County: Riv¬ erside, 22 May, 10 June 1977, D. Carroll, 2 females (UCR); Riverside, 12 May 1979, G. Cavender, 1 female (UCR); San Mateo County: San Francisco, 16 July 1963, J. and W. Ivie, 1 female (AMNH). COLO¬ RADO: Boulder County: Boulder, 1 August 1945, R. E, Gregg, 1 female (AMNH); Nederland, 16 August 1952, B. Malkin, 1 female (JCC); Denver County: Denver, 21 June 1952, R. A. Stirton, 1 female (JCC); Gilpin County: Golden Gate Canyon State Park, 28 July 2004, unknown collector, 1 male (RGH); Larimer County: Glen Haven, 27 August 1961, J. and W. Ivie, 1 male, 1 female (AMNH); Pitken County: between Aspen and Glenwood Springs along the Roaring Fork River, 12 September 1981, T. Wood, 2 males (JCC). DELAWARE: New Castle County: Wilmington, 30 June 1941, A. L. Bacon, 1 female (AMNH). DISTRICT OF COLUMBIA: Washington, September 1975, RGH, 3 males, 2 females (RGH). IOWA: Johnson County: 3 km N. Iowa City, 9 August 1984, D. K. Hoffmaster, 1 female (JCC). NEBRASKA: Scotts Bluff County: Minatare, 19 July 1978, W. F. Rapp, 2 females (JCC). NEWHAMPSHIRE: Hillsboro County: Hollis, 19 June 1987, C. D. Dondale, 1 female (CNC). NEW JERSEY: Monmouth County: Fair Haven, 30 May 1982, S. Sickerman, 1 male (JCC). NEW MEXICO: Bernalillo County: Albuquerque, 28 September 1980, J. W. Jen¬ nings, 1 male (JCC), 15 October 1982, J. W. Jennings, 1 male (JCC); Taos County: 3 km east of Taos, 6 October 1965, J. and W. Ivie, 1 female (AMNH). OREGON: numerous records of males and females from Benton COKENDOLPHER AND HOLMBERG-PHALANGIIDAE OF THE AMERICAS 11 County (UCR, ROM, OSU, TTU-Z); Douglas County (AMNH, ROM, RGH) and Lincoln County (OSU, TTU-Z); Coos County: Charleston, 3 September 1947, I. Newell, 1 female (AMNH); Curry County: Denmark, 3 August 1985, B. T. Finnamore and T. W. Thormin, 1 female (RGH); Lane County, Eugene, March 1947, B. Malkin, 2 males, 1 female (JCC); Washington County, Portland, 12 July 1971, D. J. Starkey, 1 female (JCC); Yamhill County, McMinnville, 1 June-10 July 1946, K. M. Fender, 1 female (JCC). UTAH: Cache County: Logan, 21 June 1983, 1 male (JCC); Daggett County: Red Canyon, 19 July 1949, W. J. and J. W. Gertsch, 2 males (JCC); Utah County: Provo, 3 July 1946, L. R. Allen, 1 female (AMNH). WYOMING: Park County: Bechler Ranger Station, 29 August 1982, R. Krai, 1 female (JCC). Phalangium ortoni Wood 1869 from Ecuador is a species inquirenda (Sclerosomatidae) according to Cokendolpher and Peek (1991). Phalangium pictum —see Odiellus pictus (Wood 1868). Phalangium rudipalpe Gervais 1849 is still re¬ corded in combination with Phalangium from South America (Cekalovic 1985). Because the description and illustrations (Gervais 1849) do not resemble Phalangium and because this is the only record of the family Phalangiidae south of New Mexico in the New World, it is misidentified and represents a member of some other family. From the locality (central Chile), description of the body coloration, and the illustra¬ tion of the palps, this species could be a member of Thrasychirus Simon 1884. However, the presence of the leg femora spines/tubercles is unlike that of any member of the Neopilionidae: Enantiobuninae from South America. The only other possibility is that it could be a member of the Sclerosomatidae: Gagrellinae; however, the description of P. rudipalpe is insufficient in detail to currently permit generic recognition. From a biogeographic standpoint it is best treated as a genus and species inquirenda under the Sclerosomatidae: Gagrellinae. Phalangium spinigerum Pickard-Cambridge 1905, from Guerrero, Mexico, was placed in Metopilio by Roewer (1911) (new family, “ Metopilio group”). Phalangium ventricosum = Leiobunum ventrico- sum (Wood 1868); Davis (1934) in Sclerosomatidae. Phalangium ventricosum hiemale = Leiobunum ventricosum hiemale (Weed 1890); Davis (1934) in Sclerosomatidae. Phalangium ventricosum ventricosum = Leiobu¬ num ventricosum ventricosum (Wood 1868); Davis (1934) in Sclerosomatidae. Phalangium verrucosum = Leiobunum verruco- sum (Wood 1868); Davis (1934) in Sclerosomatidae. Phalangium vittatum = Leiobunum vittatum (Say 1821); Davis (1934) in Sclerosomatidae. Subfamily Oligolophinae Banks 1893b Lacinius Thorell 1876 Two Lacinius species have been described from North America, namely Lacinius texanus Banks 1893 a and Lacinius ( Oligolophus ) ohioensis (Weed 1889a). Banks (1893a) described L. texanus from Brazos County, Texas. JCC examined the holotype (MCZ) and found it to be a penultimate female. Banks’ description is accurate but it can be added that metatarsi of leg II have 4-5 bands of setae and tibiae of leg II have no pseudosegments. There is nothing remarkable about the specimen and we conclude that it is an immature Odiellus pictus. New synonymy: Lacinius texanus Banks 1893a = Odiellus pictus (Wood 1868). We were unable to locate any other published records or collections of Lacinius (or Odiellus) from Texas, or the surrounding states of Louisiana, Arkansas, Oklahoma, or New Mexico. This especially is noteworthy as one of the oldest and largest entomological collections for Texas is located at Texas A&M University in Brazos County. Thus, we conclude that Banks’ specimen of L. texanus was probably not from Texas. Crosby and Bishop (1924) recorded “1 speci¬ men” of an unsexed L. texanus from Summerville, Georgia. In the same paper, they recorded both males and females and two other immatures as Odiellus spe¬ cies. As Bishop (1949) also recorded specimens of O. pictus from Summerville, Georgia, we conclude that Crosby’ and Bishop’s earlier report of L. texanus was an immature O. pictus. 12 Special Publications, Museum of Texas Tech University Weed (1889a) described Oligolophus ohioensis from a single “female” (later reported as a juvenile by Weed 1893) from Warren County, Ohio. Cokendolpher and Zeiders (2004) reported that this specimen appar¬ ently was lost or destroyed. Banks (1893b) transferred the species to Lacinius. In 1957, Roewer reported that he had a male (SNMF number RI/6/708) from Ohio that Banks had identified as Lacinius ohioensis , but no one has reported on the genitalia of that specimen. Walker (1928) recorded several new records of both Lacinius ohioensis and Odiellus pictus for Ohio. In her key, Walker recorded the body length of L. ohioensis as being 4-5 mm with the ocularium (= ocular tubercle) close to the margins and O. pictus as 5-6 mm long with the ocular “tubercle farther removed from the margin.” Unfortunately, she did not record the ages or sexes of the specimens examined. The specimens studied by Walker for her 1928 paper are now lost, according to one of the anonymous manuscript reviewers. Richard Bradley of the Ohio State University Chelicerates Col¬ lection (OSAL) kindly checked in their collection in April 2017 for these specimens without success. As Roewer (1923) and Bishop (1949) reported the body lengths of O. pictus as 5-6 mm, it is likely that Walker’s L. ohioensis were actually immature O. pictus. Bishop (1949) synonymized L. ohioensis with Odiellus pictus. We concur with Bishop’s synonymy. Lacinius can no longer be considered as occurring in the New World. Therefore, Lacinius species are known only from the western Palearctic (Martens 1978). Mitopus Thorell 1876 (Figs. 27-31) Mitopus morio (Fabricius 1779) (including synonyms Oligolophus montanus Banks 1893c and Opilio scabripes Walker 1860 from the New World) is patchily found throughout the northern Holarctic region (all north of about 40°N latitude) and probably has the second largest distribution of any harvestman (cf. Phalangium opilio as having the widest distribu¬ tion—although aided by humans) (Fig. 27). In North America it has been reported from northeastern areas [Kalaallit Nunaat (Greenland), Newfoundland, Nova Scotia, Quebec, Maine, New Hampshire, New York] as well as northwestern areas (Alaska, Montana/ Wyoming) (Cokendolpher and Lee 1993). The most northern records (only two above 75°N latitude) of this species are Fort Conger, Nunavut, in North America and slightly more southern on Spitsbergen Island, Norway. The Montana or Wyoming record (Savory 1933) is very unlikely as many Opiliones collections have been made from around that area without another record of this species. The color patterns of M. morio are extremely variable (see Meinertz 1973 and Martens 1978) and based solely on color it is easily confused with Phalangium opilio , which has been recorded from the region. This sample may have been incorrectly identified or labeled. This species is characterized by extreme vari¬ ability in body size, leg lengths, and body coloration and patterns. Arthofer et al. (2013) looked at Mitopus “morio ” from a single location at Tyrol, Austria, and found what they cautiously interpreted as three sym- patric cryptic species. The differences were detected with whole-genome scans, mitochondrial sequences, and morphometric characters. We agree with Arthofer et al. that much greater genetic sampling should be undertaken and hopefully will help better differentiate the various species/populations. Within North America, some western individuals have a distinct, light-colored, line running the length of the opisthosoma behind the ocularium (Fig. 28). Individuals that we have studied from eastern areas are without this pattern (Figs. 29, 30). JCC examined (in 1980) the cotypes of Oligolo¬ phus montanus Banks 1893c, and here we designate a male placed in a separate vial as the lectotype and the contents of a second vial with 2 males, 12 females, 14 immatures as paralectotypes. All of them were col¬ lected from Mount Washington, New Hampshire (Na¬ than Banks Collection, MCZ). The lectotype matches the original description of the species. The synonym Oligolophus montanus was first used in combination with Mitopus by Banks (1901). The record of M. morio as Mitopus californicus Banks 1895 from California is incorrect (see under Opilio parietinus ). Mitopus morio otherwise has not been reported from California. Except for two records that are between one and one and a half centuries old, M. morio has not been collected from the northern territories of Canada. COKENDOLPHER AND HOLMBERG-PHALANGIIDAE OF THE AMERICAS 13 Figure 27. Mitopus morio world distribution. Solid circles indicate precise locations; open circles indicate approximate locations (e.g., country records). Complete North American records from: Banks (1893c, 1900,1902); Bishop (1949); Crosby et al. (1928); Drummond et al. (2010); Hackman (1956); Koponen (1995); Lindroth and Ball (1969); Marx (1892); Meinertz (1973); Savory (1933); Walker (1860); and our collections. See text for discussion regarding Fort Conger, Port Kennedy, and Montana/Wyoming collections. Selected Eurasian and questionable African records from: Crawford and Marusik (2006); Farzalieva and Esyunin (2000); Heinajoki (1944); Hillyard and Sankey (1989); Kauri (1980); Kurt et al. (2008); Leirikh et al. (2009); Lengyel and Pall-Gergely (2010); Martens (1978); Marusik (2005); Marusik and Khrulyova (2011); Marusik and Koponen (2015); Meinertz (1973); Novak (2005); Novak et al. (2006); Roewer (1912,1924); Silhavy (1965); Slagsvold(1979); Stol (2003); Spoek(1963); Star?ga(1975,1976,1978,1984); Suzuki and Tsurusaki (1983); Tchemeris et al. (1999); and references therein. 14 Special Publications, Museum of Texas Tech University Figure 28. Preserved Mitopus morio, sex not determined, from Kenai National Wildlife Refuge, Alaska. Photograph by Matt Bowser. Figure 31. Mitopus morio , sex not determined, lateral view of chelicera, arrow points to ventral spur. Illustration traced and modified from Martens 1978. Figure 29. Mitopus morio dorsal view of female body, from New York. Illustration traced and modified from Bishop 1949. Figure 30. Mitopus morio , sex not determined, from Mont Echo, Quebec. Photograph by Pierre-Marc Brousseau. There is one record described as a new species, Opilio scabripes, by Walker (1860) from Port Kennedy, 72°N, 94°W, which would now be in the territory of Nunavut. Port Kennedy is not mentioned in any of the Canadian gazetteers or atlases that we examined. However, it is mentioned in some historical documents. William Ken¬ nedy assisted Captain Francis Leopold McClintock, and McClintock later named a harbor after Kennedy when McClintock was looking for the lost Franklin expedi¬ tion (Barr 1986). There is a tidal station, near the east end of Bellot Strait, named Port Kennedy at 72°01'N, 94°12'W. Also Fort Ross is located at 72°00'N, 94°14'W. Thus, the location recorded by Walker in 1860 is correct. The second record is from further north (Marx 1892). Marx reported that Lieutenant Greely of the Lady Lranklin Bay expedition related that Opiliones “were frequently found in the immediate neighborhood of our camp” at Lort Conger. The Tort Conger locality (now located in Quttinirpaaq National Park on Elles¬ mere Island of Nunavut, 81°45'N, 64°45'W) is about 1,200 km northwest from the nearest collection sites in Kalaallit Nunaat and represents the most northern record for the Order Opiliones (Fig. 27). The specimens from Marx have been destroyed or lost but from the location they can only be M. morio. Although it is true that Marx is notorious for his geographic mislabeling of both specimens he collected and those that came into his hands from others (Hamilton et al. 2016), COKENDOLPHER AND HOLMBERG-PHALANGIIDAE OF THE AMERICAS 15 Strand (1906) thought that this species was probably Mitopus morio ; whereas Roewer (1923) did not offer an opinion because the specimens could not be found. Cokendolpher and Lee (1993) recorded the synonymy with M. morio and we agree. Even without examin¬ ing the type material, the specimens could only be M. morio based on the original description as “blackish dingy whitish beneath; mandibles with black teeth; legs piceous, thick, comparatively short, minutely serrated,” and being collected from an inland locality at 72°N in the territory of Nunavut. As many spider collections, but no harvestmen, have been made in the far north of Canada, we are uncertain about these two old records. Port Kennedy is about 1,400 km from the closest known collection locality in Kalaallit Nunaat, 2,500 km from the closest in Labrador, and 3,200 km from the closest in Alaska. As mentioned above, the Fort Conger record is about 1,200 km from other M. morio collection sites. It is possible that individuals of this species could have been transported by the explorers to these remote locations with lumber (the sites are beyond the tree line) or veg¬ etable garden supplies (including soil) (see Lindroth 1957), or other goods. If harvestmen were transported there, they may have established local populations that by now may or may not be extinct. From the studies on cold hardiness of M. morio by Leirikh et al. (2009), cold temperature is not the limiting factor. It is also possible that some M. morio populations in eastern North America came from or were modified by early introductions by humans. New records of Mitopus morio: CANADA: NEW BRUNSWICK: St. John, 20 July 1960, E. E. Linquest, 2 females (CNC). LABRADOR: Carol Lake, n[orth] end, 13 September 1953, F. Harper, 1 female (AMNH), 12 September 1958, F. Harpen (= Harper?), 2 females (AMNH). Mitopus dorsalis Banks 1900 = Mitopus morio Fabricius (1779), new synonym. Mitopus dorsalis Banks 1900:484, plate 29, Fig. 2; Banks 1901:674; Comstock 1940:70. Nitopus dorsalis Banks 1911:420 (sic). Oligolophus dorsalis Strand 1906:474. Strandibunus dorsalis Roewer 1912:57. Parodiellus dorsalis Roewer 1923:724. “ Parodiellus ” dorsalis Cokendolpher and Lee 1993:22. Originally, Banks (1900) described Mitopus dor¬ salis based on one adult and one immature specimens collected at Popof Island during the Harriman Alaska Expedition (1899). The type specimen was deposited in the U. S. National Museum, Cat. no. 5272. The late Ralph E, Crabill (personal communication) informed JCC that the type was never deposited but more recent (2017) efforts by staff at the USNM discovered the tube with both specimens. The vial containing the two specimens had a label indicating it was “44” from the “Harriman Expedition ’99 T. Kincaid, collector” “Popoff Is 1899”. We did not separate or dissect the specimens. They are represented by two immatures, the larger (total body length 4.77 mm; femur I 3.56 mm long) we accept as the holotype and the smaller (total body length 3.73 mm; femur I 3.01 mm long) the “paratype.” The specimens now are light tan to amber colored with a very faint “clear” area where Banks indicated in his drawing of the light stripe on the opisthosoma. JCC examined a series [2 males, 1 female, 5 immatures (BMM); 1 male, 1 female, 1 imma¬ ture (NMW # 6084)] of this species that was collected at Karlyuk, Kodiak Island, Alaska, by T. Kincaid. A handwritten label in one vial indicated the collection date to be 19 July 1897, whereas other labels (typed) in the vials indicated 19 August 1897. Kincaid (1900) was in charge of the entomological collections on the Harriman Alaska Expedition. He made no mention of an 1897 collection, but did state that collections were made at Kodiak on 20 July 1899 and at Popof Island 7-17 July 1899. It appears these collection dates were mislabeled and were collected on the Harriman Alaska Expedition. The reason this series was never transmit¬ ted to Banks is unknown. The specimens from Karlyuk agree with Mitopus morio and as such are regarded as this species. These specimens are different in appearance from those from 16 Special Publications, Museum of Texas Tech University eastern North America and those from Kamkatcha. Their shorter legs and color pattern most resemble those reported from Japan by Suzuki (1939). Odiellus Roewer 1923 (Figs. 32-37) Oligolophus ohioensis = Lacinius ohioensis = Odiellus pictus. See under Lacinius for discussion of synonymy. Cokendolpher and Lee (1993) summarized the distribution of Odiellus pictus (= Phalangium pictum ) (Wood 1868) as occurring in six provinces (Manitoba, New Brunswick, both Newfoundland and Labrador, Ontario, Quebec, and Saskatchewan) and 16 states (Connecticut, Georgia, Illinois, Indiana, Maine, Mas¬ sachusetts, Michigan, New Hampshire, New York, North Carolina, Ohio, Pennsylvania, Tennessee, Ver¬ mont, Virginia, and Wisconsin). Bragg and Holmberg (2009) recorded one locality from British Columbia. Shoemaker et al. (2017) gave the first records of this species from two counties in Minnesota and we add a third county below. New records of Odiellus pictus : CANADA: ALBERTA: Baptiste Lake, 23 August-7 October 1988, RGH, 3 males (RGH); 11 km northeast of Clyde, 10-17 August 1988, T. W. Thormin, 2 males, 1 female (RGH); George Lake, 53°57’N, il4°06’W, 1 July 1973, J. Belicek, 1 female (JCC); Wagner Natural Area, west of Edmonton, 13-17 August 1985, A. T. Finnamore and T. W. Thormin, 1 male, 1 female (RGH). NOVA SCOTIA: Black River, 17 July 1956, C. D. Dondale, 1 female (CNC); Bridgewater, winter 1964-spring 1965, B. Wright, 1 male, 1 female (CNC); Coldbrook, 27 Au¬ gust 1958, C. D. Dondale, 1 female (CNC); Cow Bay, 28 September 1956, C. D. Dondale, 2 females (CNC); Halls Harbour, 22 August 1960 and 30 October 1956, 1 male and 1 female (CNC); New Grafton, 26 August 1960, C. D. Dondale, 1 male, 1 female (CNC); Prospect Bay Road, 8 October 1966, J. Gilhen, 1 male (CNC); Upper Canard, 5 September 1956, C. D. Dondale, 1 male, 2 females (CNC); Wellington, 12 October 1950, unknown collector, 2 males, 1 female (CNC). PRINCE EDWARD ISLAND: Malpaque Bay, Green Provincial Park, 5-8 September 1972, C. Starr, 2 males, 3 females (CNC). USA: MINNESOTA: Clearwater County: Lake Itasca Biological Station, Itasca State Park, August 1970, G. B. Wiggins, 4 females (ROM). Crosby and Bishop (1924) described Odiellus nubivagus from the tallest mountain peak of eastern North America in North Carolina. The type locality is Mount Mitchell at 2,037 m elevation. Nearby at the 3 rd highest peak, a new collection of what is likely this species was photographed and preserved (Fig. 32) at Clingmans Dome (2,025 m). The new specimen was not as darkly pigmented on the sides of the opistho- soma as illustrated in the original description but is considered here to represent the same species. In the original publication of the species, Crosby and Bishop (1924) reported the holotype and numerous paratypes were housed in the Cornell University and New York State Museum collections. There is now one pair of the original paratypes housed at MSU. There is a sec¬ ond sample of this species at MSU (from Grandfather Mountain) that apparently was identified by Arlan Edgar and is not a type and detailed below. The peak of Grandfather Mountain is lower at about 1,812 m. Crosby and Bishop (1924) recorded a female from Grandfather Mountain at 1,219 m elevation as being Odiellus pictus. New records of Odiellus nubivagus : USA: NORTH CAROLINA, Great Smoky Mountains National Park, Swain County, Clingmans Dome just south of parking lot, 31 July 2000, J. C. Cokendolpher, M. Hedin (1 male, GRSM 164165). Avery County, Grandfather Mountain, 4 Oct. 1960, Gertsch and Ivie, 1 male (MSU). In his dissertation, Arlan Edgar (1960) recognized Odiellus pictus as a “typical form” and as “var. I.” He presented a map (Fig. 33 on p. 58) where he showed seven counties (Chippewa, Huron, Iron, Livingston, Ontonagon, Newaygo, Tuscola) with variety 1. In 1966, Edgar described (a male and female) of his variety 1 as Odiellus pictus argentus from Michigan, without a type specimen or locality (other than state) mentioned. Later in 1971, he raised this taxon to full species and again had no types but recorded nine coun¬ ties in Michigan: adding Alger and Marquette counties. Because Edgar did not list locations in his 1966 descrip¬ tion, we are considering all of the specimens from the nine counties that still exist to be “syntypes” if they do COKENDOLPHER AND HOLMBERG-PHALANGIIDAE OF THE AMERICAS 17 Figure 32. Live Odiellus nubivagus male from Clingman’s Dome, Great Smoky Mountains National Park, Tennessee. Photograph by James C. Cokendolpher, modified; flipped horizontally so body faces left. Figure 33. Odiellus pictus male palp, ectal view. Illustration traced and modified; flipped horizontally so palp faces left, from Crosby and Bishop 1924. Figure 34. Odiellus pictus dorsal view of female body; arrow points to prosomal spine-tipped tubercles. Illustration traced and modified from Bishop 1949. Figure 35. Odiellus pictus dorsal view of male body (atypical central figure). Illustration traced and modified from Bishop 1949. Figure 36. Live Odiellus pictus male from Indian Gap, Great Smoky Mountains National Park, Tennessee. Photograph by James C. Cokendolpher. Figure 37. Live Odiellus pictus female from Athabasca, Alberta. Photograph by Robert G. Holmberg. 18 Special Publications, Museum of Texas Tech University not bear a label stating that they were collected after the date of publication in April of 1966. We have located two vials of specimens at MSU that are labeled as “ Odielluspictus argentus ” but they are not here considered to be syntypes because one vial contains specimens from outside of Michigan (and pos¬ sibly misidentified) and the other vial is not from one of the nine published counties. The non-type specimens are: 1 male, 1 female, and 1 juvenile and typewritten label “ Odiellus pictus argentus Mill Creek, P.C.334 Cheb[oygan] Co. Mich, among leaves along stream Coll. A. L. Edgar.” This sample has no date label. We did find two vials at MSU which appear to contain specimens that are “syntypes.” They are: “ Odi- elluspictus variant Probably 7th mo., Probably Huron Co.”, 9 males, 1 female (MSU) and two specimens and two labels in the second vial: “ Odiellus pictus — variant 1, Newaygo Co., Col. 7/20/57. F.R. 45.” and “Larger of 2 specimens from Livingston Co., 7/26/57 F.R. 66.” Atypewritten label from the 1960’s recorded both sets of data but is written “larger of two species” rather than “larger of two specimens.” Therefore, we are considering the smaller sized male to belong with the Newaygo County label and the female to be the Livingston County specimen. The dates correspond to dates Edgar was collecting for his dissertation so he presumably was the collector of these samples. F.R. probably means Farm Road. There are county roads running east-west through Newaygo County numbered 40, 44, and 48; but none can be located in Livingston County. With the development of these areas over the last 60 years, these farm/dirt roads may no longer ex¬ ist. These latter two specimens are here designated the lectotype (male Newaygo County) and paralectotype (female Livingston County). We have separated the two specimens and re-labeled the vials so no confusion will exist in future studies. We also designate the Huron County specimens all as paralectotypes. The lecto¬ type male and paralectotype female (from Livingston County) match the original description of the species (except much faded in color—white to silver, tinged with pink is now creamy to light tan in color), including the measurements. The “white spines” on the abdomen are little more than granules that are spine-tipped, not very noticeable. The total length of the lectotype was taken from the anterior-most point of the prosoma to the end of the opisthosoma. This anterior point is not in the center in front of the trident, but an imaginary line drawn from the forward pointing sides of the prosoma in front of each ozopore. The paralectotypes from Huron County match the description, but are slightly smaller in measurements. Neither O. argentus nor O. nubivagus has been recorded outside the states in which they were origi¬ nally described. As O. pictus is such a widespread and variable species and O. argentus and O. nubivagus seem to be restricted to very local areas, it appears that this genus needs a thorough taxonomic study of North American specimens to determine the relationships. The penis has only been illustrated by Bishop (1949: Figs. 27,28); O. pictus specimen presumably from New York. We have used a very conservative approach by identifying many collections as O. pictus. Oligolophus C. L. Koch 1871 (Figs. 38, 39) Oligolophus tridens (C. L. Koch 1836) is known from Iceland, northern through central Europe, and was first reported in America as being introduced into Vermont prior to 1974 (Bell 1975). However, Hack- man (1956) speculated that his immature Oligolophus sp. from Newfoundland was this species. Bragg and Holmberg (2009) gave their earliest collection dates for British Columbia, New Brunswick, Newfoundland, Nova Scotia, Ontario, Prince Edward Island, Quebec, and Maine. Our work indicates that this species was present in Newfoundland as early as 1947 [i.e., St Figure 38. Live Oligolophus tridens , sex not determined, from the Netherlands. Photograph by Jan van Duinen. COKENDOLPHER AND HOLMBERG-PHALANGIIDAE OF THE AMERICAS 19 Figure 39. Oligolophus tridens male palp, ectal view. Illustration traced and modified; flipped horizontally so palp faces left, from Martens 1978. John’s, 27 October 1947, R. Traub, 1 male, 2 females (JCC)]. All records are from introductions into eastern and western North America. Oligolophus ohioensis = Odiellus pictus; see Lacinius for details. Oligolophus montanus = Mitopus morio\ see Mitopus for details. Par oligolophus Lohmander 1945 (Figs. 40—43) Bragg and Holmberg (1975) reported that Par- oligolophus agrestis (Meade 1855) was introduced into British Columbia and Washington. The species is otherwise known from Europe. Bragg and Holmberg (2009) recorded new records from British Columbia as well as noting collections from Alberta and Nova Scotia. The earliest east coast record [Nova Scotia: Halifax, 29 September 1950, unknown collector, 1 female (CNC)] coincides with the earliest west coast record that we have found [i.e., British Columbia: Vancouver Island, Wellington, 20-24 August 1950, R. Guppy, 1 female (JCC)]. Figure 40. Par oligolophus agrestis female genital operculum with a large notch on anterior border. Illustration traced from Martens 1978. Figure 41. Paroligolophus agrestis male genital operculum with a small notch on anterior border. Illustration traced from Martens 1978. Figure 42. Paroligolophus agrestis penis. Illustration traced from Martens 1978. Figure 43. Live Paroligolophus agrestis , sex not determined, from Spanaway, Pierce County, Washington. Photograph by Joseph V. Higbee. 20 Special Publications, Museum of Texas Tech University Subfamily Opilioninae C. L. Koch 1839a Egaenus Koch, in Hahn and Koch 1839b The only species from the New World described in this otherwise Eurasian genus is Egaenus mexicanus Becker 1886. No specific locality in Mexico was re¬ corded by Becker. Currently, the species is considered a synonym of Opilio ischionotatus Duges 1884 (= Leiobunum ischionotatum ), see below under Opilio. Egaenus is not known from the Americas. Opilio Herbst 1798 (Figs. 44, 45) Because of the antiquity of the name, some of the older specific names for New World harvestmen have been associated incorrectly with Opilio. Roewer (1952, 1956) described three new species from the New World under Opilio. All have been moved to other genera or synonymized. In 1955, Levi and Levi indicated (in a footnote) that Opilio angulatichelis Roewer 1952 from “Glacial” (Glacier) National Park, Montana, was an immature male Phalangium opilio. JCC examined the holotype (SNMF, vial RII/11048/328 with body and legs; micro¬ scope slide number 16107 with left palp and chelicera) and found it to be as Levi and Levi suggested (Coken- dolpher and Lee 1993). The smooth supracheliceral laminae and reduced spur on the chelicerae are normal for immature male P. opilio. Opilio bolivianus Roewer 1956 was synonymized under Cristina bispinifrons Roewer 1917 (subfamily Phalangiinae) by Cokendolpher and Rylander (1986). However, they concluded that the collection locality of Bolivia was incorrect and suggested that it originated from the Cameroon region of western Africa. Opilio chickeringi Roewer 1956, from Costa Rica, became the type species for the genus Lanthan- opilio Cokendolpher and Cokendolpher 1984. This species was first placed in the Phalangiinae by Cok¬ endolpher and Cokendolpher (1984) but provisionally was transferred to the Dicranopalpus genus group (family incertae sedis) by Crawford (1992). Both Opilio gertschi and Opilio mexicanus Roewer 1956, from Mexico (Tamaulipas), were transferred to the Figure 44. Live Opilio parietinus male from Baptiste Lake, Athabasca County, Alberta. Photograph by Robert G. Holmberg. Figure 45. Live Opilio parietinus female from Baptiste Lake, Athabasca County, Alberta. Photograph by Robert G. Holmberg. genus Metopilio Roewer 1911 (new family, “ Metopilio group”) by Cokendolpher and Cokendolpher (1984). We also recognize Lanthanopilio as being a member of that same new family. The reason this genus was first placed in the Phalangiidae was based on the morphol¬ ogy of the penis: bent junction of the glans and shaft. This derived condition apparently was developed in¬ dependently a second time in the “Metopilio group.” The biogeography and other morphological characters place this species in the “Metopilio group.” Opilio scabripes Walker 1860 = Mitopus morio ; see Mitopus for details. COKENDOLPHER AND HOLMBERG-PHALANGIIDAE OF THE AMERICAS 21 Mitopus californicus Banks 1895 = Opilio pari- etinus (De Geer 1778), new synonym. Mitopus californicus Banks 1895:66, 1901:674, 1904:362; Myers 1921:19; Comstock 1940:69. Nitopus californicus Banks 1911:420 ( lapsus calami). There are two cotypes of Mitopus californicus in the Nathan Banks collection (MCZ). The male is hereby designated the lectotype, the female a paralec- totype. Examination of the types reveals that they are both specimens of Opilio parietinus (De Geer 1778) and not Mitopus morio as indicated by Roewer (1912) and subsequent authors. The type specimens of M. californicus were reported (Banks 1895) to have been from Los Angeles, California. Like so many other harvestmen described by Banks from southern Cali¬ fornia, the type locality data appear to be in error. In the USA, Opilio parietinus is only encountered in the more northern regions; no valid records from southern California are known. The above discussion leaves Opilio parietinus (De Geer 1778) as the only Opilio in the New World. It is widely distributed in the western Palearctic and introduced into Tasmania (Martens 1978). It was first described from North America from New York as Pha- langium cinereum Wood 1868. Human transport was possibly responsible for aiding the movement of this species around the globe (Gruber and Hunt 1973) and it was likely introduced into North America from Europe (Bragg and Holmberg 2009). It is uncommon in col¬ lections from the Nearctic and seldom occurs in great numbers as Phalangium opilio , with which it is often collected and sometimes confused. Cokendolpher and Lee (1993) recorded this species from three Canadian provinces (Ontario, Quebec, and Saskatchewan) and 18, mostly northern, states (Colorado, Idaho, Illinois, Iowa, Maine, Massachusetts, Michigan, Montana, Nebraska, New Hampshire, New Mexico, New York, North Dakota, Ohio, Pennsylvania, South Dakota, Texas, Vermont, and Wisconsin). Bragg and Holmberg (2009) noted records from British Columbia as well as Alberta and Manitoba. Additional state records are: USA: UTAH: Box Elder County: Raft River Mountains, Lynn, “9-8-32”, R. V. Chamberlin and W. Ivie, 3 males, 4 females (AMNH); Carbon County: Price, November 1942, H. and M. Higgins, 2 males, 3 females (AMNH); Daggett County: Red Canyon, 19 July 1949, W. J. and J. W. Gertsch, 1 male (JCC); Garfield County: Henry Mountains, Horse Valley, “9-6-29”, R. V. Chamberlin, 1 female (AMNH); Salt Lake County: Mill Creek Can¬ yon, no date, R. V. Chamberlin, 1 female (AMNH); Salt Lake City, 18 October 1930, W. Ivie, 2 males, 1 female (AMNH); Salt Lake City, 30 October 1956, R. Rosella, 2 females (AMNH); west of Salt Lake City, no date, H. Fowler, 1 female (AMNH); unknown county: specific locality?, 14 May 1931, Yu-hsi M. Wang, 4 males, 9 females (AMNH); WYOMING: unknown county: “So. Mountainea”, “8-6-46”, collector unknown, 1 female (AMNH). Opilio ischionotatus Duges 1884 (= Egaenus mexicanus Becker 1886) from Guanajuato and Michoa- can, Mexico, is now known (Kury and Cokendolpher 2000) in the combination Leiobunum ischionotatum (Sclerosomatidae: Leiobuninae). Platybuninae Star^ga 1976 Lophopilio Hadzi 1931 (Fig. 46) The single species of the genus recorded from the region is Lophopilio palpinalis (Herbst 1799) from Canada. This species is otherwise known from across Central and Northern Europe (Martens 1978) and Latvia, Lithuania, Byelorussia, Ukraine, Russia: Figure 46. Live Lophopilio palpinalis , sex not determined, from the Netherlands. Photograph by Jan van Duinen. 22 Special Publications, Museum of Texas Tech University Kaliningrad, Leningrad and Moscow areas, Karelia (Farzalieva and Esyunin 2000). Although this intro¬ duced species resembles the native Odiellus pictus , they are easily told apart by the absence of a hook on the ventral surface of the basal cheliceral segment in Lophopiiio. This is a good character as it can be seen on adults as well as immatures. New record of Lophopiiio palpinalis : CANADA: NEW BRUNSWICK: Fundy, Laverty Falls, 11 August 2009, J. Cossey, J. Fernandez, R. Labbee, J. Smith, M. Zhang Mixed forest, N45.6570015, W65.01499939, 404 m. elev., 1 male (Biodiversity Institute of Ontario, University of Guelph). Megabunus Meade 1855 (Fig. 47) Megabunus diadema (Fabricius 1779) is a spe¬ cies of harvestman widely distributed in Europe, where it has been found in Iceland, Faroe Islands, western Norway, Great Britain, western France, Belgium, and northern Spain (Stol 2005). It presumably was intro¬ duced into Iceland and possibly will be introduced to other areas west with shipping and commerce in the Americas. This species was reported as being mainly parthenogenetic by Stol (2005), which would greatly increase the odds in establishing a population with only a single specimen. More recently populations that are not parthenogenetic have been discovered (D’Amico Figure 47. Preserved Megabunus diadema female from England. Photograph by Robert G. Holmberg. and Danflous 2015) but they are in the warmer/southem reaches of the distribution in Spain. Rilaena Silhavy 1965 (Figs. 48-51) Martens (1978) discussed European species of Rilaena. Bragg and Holmberg (1975) reported that Rilaena triangularis (Herbst 1799) was introduced into British Columbia and Washington. Bragg and Holmberg (2009) noted that the earliest North American record was 1951 (Washington) but there also were col¬ lections from Maine (see also Drummond et al. 2010), Massachusetts, and New York. The first record from Quebec was photographs by Pierre-Marc Brousseau Figure 48. Live Rilaena triangularis subadult male from Kentville, Nova Scotia. Photograph by Robert G. Holmberg, modified; flipped horizontally so body faces left. Figure 49. Rilaena triangularis male from Mont St-Bmno, Quebec. Photograph by Pierre-Marc Brousseau. COKENDOLPHER AND HOLMBERG-PhALANGIIDAE OF THE AMERICAS 23 Figure 50. Rilaena triangularis anterior view of prosoma and chelicerae (2nd segment with large distal spur) of male from Mont St-Bruno, Quebec. Photograph by Pierre-Marc Brousseau. Figure 51. Rilaena triangularis lateral view of chelicera, arrow points to region where ventral spur is missing, sex not determined. Illustration traced and modified from Martens 1978. (Figs. 49, 50 herein) on the internet (https://bugguide. net/node/view/897539/bgimage). Some other au¬ thors have reported triangularis in combination with Paraplatybunus Dumitrescu 1970 and Platybunus C. L. Koch 1839a, but, most recent authors do not. We herein report R. triangularis for the first time from Nova Scotia (Fig. 48). New records of Rilaena triangularis : CANADA: NOVA SCOTIA: Cape Breton Island, St. Anns, 22 June 2005, RGH, 1 subadult male (RGH); Kentville, 24 June 2005, RGH, 15 subadults, 3 of which have the cheliceral spurs typical of males (RGH). Misplaced Species Protolophidae Banks 1893b Parodiellus Roewer 1923 The single species from the Americas recorded in Parodiellus is misplaced in that genus. Cokendol- pher and Lee (1993) indicated that this species may have been placed in the wrong family. The species correctly is placed in Protolophus Banks 1893b of the Protolophidae Banks 1893b. Parodiellusprojectus Goodnight and Goodnight 1942b (Phalangiidae) = Protolophus niger Goodnight and Goodnight 1942a (Protolophidae), new synonym. Goodnight and Goodnight (1942b) described Parodiellus projectus from Oregon. Examination of the male (penultimate) holotype (by JCC) revealed that it is conspecific with Protolophus niger Goodnight and Goodnight 1942a. As the name P. niger was published 22 January 1942a, and P. projectus was published 13 February 1942b, the later name is the junior synonym. The holotype of P. projectus apparently was collected with part of the type series of P. niger at Rogue River Valley, Oregon (April 1934, Lawrence). It presumably was considered different due to the fact that it was in the process of the final molt when it was preserved and therefore the external morphology was differ¬ ent. Adult members of the genus Protolophus Banks show considerable sexual dimorphism (Goodnight and Goodnight 1942a; JCC personal observation); whereas penultimate males show characteristics of both sexes. Fortunately, the holotype of P. projectus has a well-formed penis making identification certain. The holotype of P. projectus and the type series of P. niger are deposited in AMNH. 24 Special Publications, Museum of Texas Tech University Taxonomic Key to the Phalangiidae of the Americas The following key primarily is based on Edgar’s publication (1990). The key is applicable for adults, most subadults, and older immatures. For positive identifications, mature males are best. Females, sub¬ adults, and older immatures often need to be associated with mature males for positive identification. Most immatures cannot be identified to more than family level unless one already knows what species occur in a specific area and has an extensive knowledge about their variability. Any key only provides tentative identification, therefore, one must also check other characteristics pro¬ vided in Table 1 and the text as well as in the literature cited. To confirm identification of European species, see Hillyard and Sankey (1989), Martens (1978), Spoek (1963), and Uddstrom and Rinne (2016). The latter has excellent color photographs of most European species covered here. For Leptobunus species, see Cokendol- pher (1985). For Liopilio species, see Cokendolpher (1981a). Bishop (1949) still has the best illustrations for most eastern species. The best morphological and anatomical charac¬ teristics (in descending order) are on the: penis [consist¬ ing of a long, basal shaft (corpus) topped with a bulbous glans that has a long, terminal spine-like stylus; in lateral view, the glans is bent about 90 degrees from the corpus]; palp (= pedipalp, palpus); spine-tipped tuber¬ cles on the anterior dorsal prosoma (= cephalothorax); chelicera; ocularium (= ocular tubercle, eye turret); genital operculum; coloration; other characteristics. The segments of the palp, from the base, are as follows: coxa [sometimes with an apophysis (= lobe; distal end expanded on inner/mesal margins) and/or spine-tipped tubercles], trochanter (sometimes with ventral spine- tipped tubercles), femur (longest segment; sometimes with ventral, elongate spine-tipped tubercles), patella (usually with a small to large mesal apophysis), tibia (usually with a small to large mesal apophysis), and tarsus (males may have a ventral series of denticles; ensure the claw is smooth, not toothed). The sizes of bodies and body parts are variable. For specimens preserved in alcohol, one can determine maturity by examining the anterior edge of the genital operculum. In adult specimens, the anterior edge is open and can be lifted slightly with fine-tipped forceps or a needle. With forceps, the genital oper¬ culum can be pulled/ripped backwards to reveal the penis or ovipositor. In immatures, the anterior edge of the operculum is fused to the body and the penis or ovipositor is absent or only partially developed. Phalangiids are drably colored (shades of brown, grey, black, tan, yellowish and white). The pigmenta¬ tion patterns can be useful for identification but are inconstant because of genetic variability as well as sex, age, and stage within the molting cycle. Generally, adult males are more darkly pigmented, more armored and smaller, than females. Methods of preservation and lengths of time of preservation also affect color. Colors referred to in the key apply to specimens preserved in alcohol. To observe the seminal receptacles, the tip of the ovipositor must be cleared (clove oil or lactophenol work well; see Acosta et al. 2007). COKENDOLPHER AND HOLMBERG-PhALANGIIDAE OF THE AMERICAS 25 1 . r. 2 . 2\ 3. 3’. 4. 4\ 5. 5’. 6 . 6\ 7. T. 8 . 8 ’. 9. 9’. Key Anterior median border of prosoma with one to three conspicuous spine-tipped tubercles much longer than any other tubercles in the anterior area (area in front of the eyes); the tubercles usually form a trident with the central tubercle the longest (Figs. 34, 35, 46, 47).2 Anterior median border of prosoma smooth, granulated, or with one to several small spines/tubercles of near equal length (Figs. 10, 16, 17, 22, 38).8 Ocularium spines/tubercles not exceedingly prominent, shorter than the eye diameter.3 Ocularium spines/tubercles longer than the eye diameter and nearly the height of the ocularium (Fig. 47) (in Europe and Iceland, not yet found in mainland North America). Megabunus diadema Genital operculum with a smooth, rounded anterior margin; ocularium with two longitudinal rows of small tubercles.4 Genital operculum with a medial notch on the anterior border (Figs. 40, 41); ocularium nearly smooth with a few setae only (Fig. 43). See also Fig. 42. Paroligolophus agrestis Palpal femur with several large spine-tipped tubercles ventrally (Fig. 33).5 Palpal femur with only spines or a few short spine-tipped tubercles (Fig. 39). See also Fig. 39. . Oligolophus tridens Basal segment of the chelicera with a ventral hook (spur) (like in Fig. 31). Odiellus .6 Basal segment of chelicerae smooth, without a ventral hook (like in Fig. 51). See also Fig. 46. . Lophopilio palpinalis Central figure (dorsal, bilaterally symmetrical, pigmented pattern on body; also called a saddle) distinct or not, generally more obscure in females, mottled with brown, remaining dorsum silvery with brown wisps (Figs. 34-37); femur of leg II tan but blotched with dark brown and ringed with brown distally; ocularium generally lighter except beneath the eyes. Odiellus pictus Central figure of dorsum not conspicuously mottled but uniform dark brown or black, except for a few small white spots (Fig. 32); femur of leg II tan to black with little or no distal dark ring; ocularium dark, not contrasting with the central figure.7 Lateral borders of central figure distinct, nearly parallel on opisthosoma (abdomen), dark brown, remaining dorsum silvery white, female with tan and pink tinge, diverging anteriolaterally on prosoma; femur of leg II tan with little or no distal dark ring; ocularium a bit lighter than central figure; from Michigan. . Odiellus argentus Legs, sides of opisthosoma and much of prosoma including ocularium dark brown to black; from high mountains of North Carolina and Tennessee (Fig. 32). Odiellus nubivagus Ocularium with spine-tipped tubercles.14 Ocularium smooth or, at most, small setae.9 Palpal femur and patella expanded mesally (Figs. 14, 15); Male: palpal tarsus ventrally with denticles (Fig. 14) . Liopilio .13 Palpal tibia and tarsus may be expanded mesally (Figs. 1,11), femur and patella not expanded mesally (Fig. 2); Male: palpal tarsus ventrally without denticles. Leptobunus .10 Special Publications, Museum of Texas Tech University 26 10. Male: distal segment of chelicera with distinct lateral spur (apophysis) on movable finger (Fig. 3); stylus of penis nearly equal to or shorter than setae of glans (Figs. 4, 5); Female: primary loop of seminal receptacle short, less than three times as long as wide (Fig. 6). Slimlepto new subgenus.11 10’. Male: distal segment of chelicera without lateral spur on movable finger; stylus of penis much longer than setae of glans (Figs. 7, 12); Female: primary loop of seminal receptacle long, more than four times as long as wide (Figs. 8, 9). Leptobunus subgenus.12 11. Body color usually golden; legs speckled; Male: setae of glans separated by more than the length of the stylus (Fig. 4); only known from California. Leptobunus aureus XV. Body color brown, gray, or bronze; legs banded; Male: setae of glans closely spaced (Fig. 5). Distributed from Alaska to California. See also Fig. 10. Leptobunus parvulus 12. Male: palpal tibiae and tarsi expanded on mesal side (Fig. 1); dorsum of penis glans slightly depressed (Fig. 7); Female: basal loops of seminal receptacles consisting of three coils or loops (Fig. 8). . Leptobunus californicus XT. Male: palpal tibiae and tarsi only slightly expanded (Fig. 11); dorsum of penis glans straight (Fig. 12); Female: basal loops of seminal receptacles consisting of two coils or loops (Fig. 9). See also Fig. 13. . Leptobunus borealis 13. Central figure on opisthosoma distinct and not cross-shaped (Fig. 16); body sometimes yellowish; Male: palpal patella and tibia armed ventrally with small spines as well as denticles (Fig. 14). See also Fig. 15. Liopilio yukon 13’. Central figure on opisthosoma obscured, with light and dark spotting, or absent (Figs. 17-19); body often yellowish green; Male: palpal patella and tibia without small spines but with denticles. See also Figs. 20-21. Liopilio glaber 14. Basal segment of chelicera ventrally with a sharp, forward-pointed hook (spur) (Fig. 31). See also Figs. 28-30. Mitopus morio 14’. Basal segment of chelicera ventrally smooth, without a hook (like in Fig. 51).15 15. Superacheliceral laminae smooth; Male: chelicerae and palps normal.16 15’. Supracheliceral laminae, of adults and subadults, with two anteriorly directed spines visible below anterior margin of prosoma (Figs. 22, 24); Male: distal segment of chelicera elongated dorsally as a spur (length is variable) and palps very slender and elongate, extending considerably above the prosoma (Figs. 23, 25); Female: see Fig. 26. Phalangium opilio 16. Ocularium strongly canaliculate (grooved) and about one-third the width of the prosoma (Figs. 49, 50). Palp with a distinct apophysis on the inner side of the patella and a much smaller one on the tibia (Figs. 48, 49). Male with large spur on 2 nd segment of chelicera (Figs. 50, 51). Rilaena triangularis 16’. Ocularium not canaliculate and about one-fifth the width of the prosoma (Figs. 44-45). Palpal segments nearly cylindrical with only slight apophyses. Male femur I thicker than that of other legs and all female legs (Figs. 44, 45). Opilioparietinus COKENDOLPHER AND HOLMBERG-PHALANGIIDAE OF THE AMERICAS 27 3 • CD O C x Oh o i B o C/3 PP E^ a cd CD Oh o o a, 3 Oh Oh C/3 s = I to P S cd O O cd ’ H o pp X h-> co n S 3 O W C 3 CD CD CD d -0 cn "o Oh H O 3 3 033 3 u ^ Oh ^ £ P Pd 3 .tn CO ;> 3 d ^ d 3 o 5b E^ I S o V! 3 033 3 CD - . P 3 n o u P- 2 Oh .5 3 ' t_ ‘ O Oh 3 PP O r-r w w g U O ^ p CD 2 O cd CL P CN co co s_L *—i r\ /is . 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Typical with Central figure Reported from ate, tapering patella with smooth, devoid with or without Dark between smooth distal distinct, widest Alaska and distally with a prominent of spines. Fig. ventral hook. the eyes. Fig. margin. at tergite 3 and Yukon. COKENDOLPHER AND HOLMBERG-PHALANGIIDAE OF THE AMERICAS 31 :— 1 2 vri ^ CO '&.SP vd ° Ph CD 45 ■ti .22 g £P ^ £ H o o CO CO a © 73 _g 13 _ 03 d ^ g T3 S C1J ^ T3 73 —h C • • ® B) g .2 U g s s z a ..AM 8 s .£» 13 'S Ph c « k -g -3 "S C/2 t: o ^ o Ph 2 Dh g •a 2 -s a -a b p § g CD O CD g -E3 •§ £ 3 ° g CO • S tS £ < o o ^ 'S ^ Gh 1/13 C3 2~i ■•—> CD .9r Ph rri -4—* CD c /2 g g ^ &, g .b g i g § a a^j C/2 Jg C/2 C/2 fN CD . O m ^ 2 o *2 ^ g g c ^ K S ^ T3 t: _ O CD 4C CD C/2 C 2 ° £ g ’d 3 8.1 a £ co aS ' I D g ^ o a* 1*1 D y. cw Oh Ji o 13 CD g D >? b o .b 43 ^ bb -g g -tb D D2 T3 bb £ . « £ .§> § vd Ph Ph (N CD O CD g D 3 CD g g '-O O +3 g g ^ o g 22 U g T3 i 81 s ° D CZJ C/2 > g g £ £ ^ * 2 13 2 E E> S 3 03 o oj a -a o > PP > jo --0 o> aa u T3 ■“ 03 3 Oh aa So o O 03 O T3 O _ ‘C 03 E -O < 'I I E 03 03 T3 Oh E Oh VO 133 i e s £ Oh 03 O cn 'Sh y o „ E j§ -S ta ~ 13 e a ^ 6 aa o _o E a . tas E E g 'P 03 fi 1 * IN a •■i o ab E 133 03 ^ ca O j_p g ‘g >b | i £ -S .2 & • 03 T3 O E ca ^ -O u -S CLI » HZ g 1 3 S t 133 133 133 03 o O 5 ca JS E O §<§ o' Q ^ cx, a E 1 3 a a js m ^ r ■> T3 03 03 03 c3 03 tS ^ . . E _ O >» cl G o* Oh 'S c O 2 03 a o o E ts E 13 W -O a •tn 133 ^ "3 B E .y © Oh O E 1 ® M oo f £ a o .o 03 s ! 1. 1 1 § I “ IJ - s i $ o a o ^ a J) M n in -S 5 W 5 5> E ta a C t-, PP ^ 4=1 E ca J-H i-H 03 a < 03 03 03 O -9 ^ .a c 9 ■' 03 "2 E aa Oh +h ab E 03 £ -3 % M l> - 03 m & 03 03 .£ o -I 03 3 ao 33 o a “* rl & ^ W --0 o s = .s y o = £ E ao ca 03 Oh ' E JS 9 .23 £ T3 (03 03 u E -S < -B Oh Oh O o Oh ca = 1 ! •§ Ph a aa E 133 03 33 >3 •a Oh N ca tj- &* u 5 a I a -S S Sb ^3 a g E O B E a ca E 03 S 3 ^ aa E -9 o £*•> a 03 ;> a B C33 c3 03 O Genital Species Penis Palp Prosoma Chelicera Ocularium Operculum Coloration Comments Rilaena triangu- Corpus slender Moderately Anterior area Basal segment 2 rows of 7 to Typical with Central figure Now found laris but with a broad large spine- only with a few without ventral 11 small spine- smooth distal variable; usually in eastern and base. Gians tipped tubercles small spine- hook. Male: tipped tubercles, margin. brown, spotted western North COKENDOLPHER AND HOLMBERG-P HAL AN GIID AE OF THE AMERICAS 33 c—> o B d d P <3 "p a § < "O E o' d tax) S 1 o a> d ^3 . _h J-h p Mh p p p £ 03 3 P o £ 3 Ph v 3 p "d gsl = !§'=!„ o £ ^ £ -o SO -d 3 o 3 £ o d tax) 8 E £ , 3 PLh °' P LO p tap E p a & p 5 tax) -a — ^ c d _ —- ' r~i Cu S s ^ ^ .2 O 3 o" 5 i3 a o in p ^ £ d £ p 03 ^ : I1 d "I gP £ 2 T3 -5 o fl) ^ ^ £ . d o a ^ s l ' i—t Os - „ p d 3 P d 1/2 d cn £ d P S tax) E x d cn ^ O 3 g 34 Special Publications, Museum of Texas Tech University Summary The conclusions of this study are summarized in Table 2. Of the 68 scientific names considered, only 16 species of Phalangiidae in 10 genera are still valid for the Americas. No Phalangiidae are known to occur in Mexico, the Caribbean, Central or South America. Egaenus and Lacinius do not occur in the Americas. The single species of each of the genera Lophopilio, Oligolophus , Opilio , Paroligolophus, Phalangium, and Rilaena were introduced accidentally by humans from Europe to more northern locations of North America. It is unclear if Mitopus morio is native to parts of North America and introduced to others; it is now found in the northeastern and northwestern areas of the conti¬ nent. A taxonomic key to the members proper of the Phalangiidae is presented. Table 2. Summary of the Family Phalangiidae in North America, listed alphabetically. There are no true Phalangiidae species in Mexico, Central and South America, or the Caribbean. Species Taxonomic Status Distribution in North America Native / Introduced Comments Egaenus mexicanus = Leiobunum ischiono- tatum — — Sclerosomatidae: Leiobun- inae Lacinius ohioensis - Odiellus pictus — — Phalangiidae: Oligoloph- inae Lacinius texanus = Odiellus pictus — — Phalangiidae: Oligoloph- inae, new synonymy Leptobunus atavus = Amauropilio atavus Fossil - Colorado — Sclerosomatidae: Leiobun- inae Leptobunus aureus Valid California Native Phalangiidae: Phalangiinae Leptobunus borealis Valid Alaska and nearby Russian islands Native Phalangiidae: Phalangiinae Leptobunus californicus Valid California and Nevada Native Phalangiidae: Phalangiinae Leptobunus mexicanus = Paranelima mexicana — — Sclerosomatidae: Leiobun- inae Leptobunus parvulus Valid West coast, Alaska to California Native Phalangiidae: Phalangiinae Leptobunus spinulatus = Metopilio spinulatus — — “ Metopilio group”, not Phalangiidae Leuronychus parvulus = Leptobunus parvulus — — Phalangiidae: Phalangiinae Liobunum parvulum = Leptobunus parvulus — — Phalangiidae: Phalangiinae Liomitopus laevis = Leptobunus californicus — — Phalangiidae: Phalangiinae Liopilio glaber Valid Rocky Mountains, south of L. yukon Native Phalangiidae: Phalangi¬ inae, new record from 1 state Liopilio yukon Valid Alaska, Yukon Native Phalangiidae: Phalangiinae Lophopilio palpinalis Valid New Brunswick Introduced from Europe by 2009 Phalangiidae: Platybun- inae, new record from 1 province COKENDOLPHER AND HOLMBERG-PHALANGIIDAE OF THE AMERICAS 35 rable 2. (cont.) Species Taxonomic Status Distribution in North America Native / Introduced Comments Megabumis diadema Valid European spe¬ cies introduced to Iceland and not yet found in North America Phalangiidae: Platybuninae Mitopus californicus = Opilio parietinus — — Phalangiidae: Opilioninae, new synonymy Mitopus dorsalis = Mitopus morio — — Phalangiidae: Oligoloph- inae, new synonymy Mitopus morio Valid Northwest and north¬ east North America, including Kalaallit Nunaat Native but pos¬ sibly introduced in some areas in the northeast and Alaska Phalangiidae: Oligolo- phinae, new records for 2 provinces Nitopus dorsalis = Mitopus morio — — Phalangiidae: Oligoloph- inae, lapsus, misspelling of genus Odiellus argentus Valid Michigan Native Phalangiidae: Oligoloph- inae Odiellus nubivagus Valid North Carolina, Ten¬ nessee Native Phalangiidae: Oligoloph- inae Odiellus pictus Valid Cool, wet forests across southeastern and northern North America Native Phalangiidae: Oligoloph- inae, new records from 3 provinces and 1 county Oligolophus dorsalis = Mitopus morio — — Phalangiidae: Oligoloph- inae Oligolophus montanus = Mitopus morio — — Phalangiidae: Oligoloph- inae Oligolophus ohioensis — Odiellus pictus — — Phalangiidae: Oligoloph- inae Oligolophus tridens Valid Eastern North America and British Columbia Introduced from Europe by 1947 Phalangiidae: Oligoloph- inae Opilio angulatichelis = Phalangium opilio — — Phalangiidae: Phalangiinae Opilio bolivianus - Cristina bispinifrons West Africa, not South America — Phalangiidae: Phalangiinae Opilio chickeringi = Lanthanopilio chick¬ eringi — — “ Metopilio group”, new status Opilio gertschi = Metopilio gertschi — — “ Metopilio group” Opilio ischionotatus = Leiobunum ischiono- tatum — — Sclerosomatidae: Leiobun- inae Opilio mexicanus = Metopilio mexicanus “ Metopilio group” 36 Special Publications, Museum of Texas Tech University Table 2. (cont.) Distribution in Native / Species Taxonomic Status North America Introduced Comments Opilio parietinus Valid Northern U.S.A. and southern Canada Possibly in¬ troduced from Europe by 1868 Phalangiidae: Opilioninae, new records from 2 states Opilio scabripes — Mitopus morio — — Phalangiidae: Oligoloph- inae Paraplatybunus triangu¬ laris = Rilaena triangularis — — Phalangiidae: Platybuninae Parodiellus dorsalis = Mitopus morio — — Phalangiidae: Oligoloph- inae Parodiellus projectus = Protolophus niger — — Protolophidae, Protoloph- inae, new synonymy, new family status “ Parodiellus ” dorsalis = Mitopus morio — — Phalangiidae: Oligoloph- inae Parol igolophus agrestis Valid Eastern and western North America Introduced from Europe by 1950 Phalangiidae: Oligoloph- inae Phalangium acanthipes = Metopilio acanthipes — — “Metopilio group” Phalangium armigerum = Metopilio armigerus — — “Metopilio group” Phalangium bicolor = Nelima elegans — — Sclerosomatidae: Leiobun- inae Phalangium calcar = Leiobunum calcar — — Sclerosomatidae: Leiobun- inae Phalangium cinereum = Opilio parietinus — — Phalangiidae: Opilioninae Phalangium dorsatum - Leiobunum vittatum — — Sclerosomatidae: Leiobun- inae Phalangium exilipes = Leiobunum exilipes — — Sclerosomatidae: Leiobun- inae Phalangium favosum = Trachyrhinus favosus — — Sclerosomatidae: Gagrel- linae Phalangium formosum = Leiobunum formosum — — Sclerosomatidae: Leiobun- inae Phalangium grande = Hadrobununs grandis — — Sclerosomatidae: Leiobun- inae Phalangium horridum = Metopilio cambridgei — — “Metopilio group” Phalangium longipalpus = Phalangium opilio — — Phalangiidae: Phalangiinae Phalangium maculatipes = Metopilio maculatipes — — “Metopilio group” Phalangium maculosum = Hadrobunus macu- losus — — Sclerosomatidae: Leiobun- inae Phalangium nigropalpi — Leiobunum nigropalpi — — Sclerosomatidae: Leiobun- inae COKENDOLPHER AND HOLMBERG-PHALANGIIDAE OF THE AMERICAS 37 Table 2. (cont.) Species Taxonomic Status Distribution in North America Native / Introduced Comments Phalangium nigrum — Eumesosoma nigrum — t — Sclerosomatidae: Leiobun- inae Phalangium opilio Valid Southern Canada and northern U.S.A. Possibly in¬ troduced from Europe by 1890 Phalangiidae: Phalangi- inae, new records from 19 provinces/states Phalangium ortoni species inquirenda — — Sclerosomatidae: Gagrel- linae Phalangium pictum = Odiellus pictus — — Phalangiidae: Oligoloph- inae Phalangium rudipalpe Genus and species inqui¬ renda — — Sclerosomatidae: Gagrelli- nae, new status; described from central Chile Phalangium spinigerum = Metopilio spinigerus — — “ Metopilio group” Phalangium ventricosum hiemale — Leiobunum ventrico¬ sum hiemale — — Sclerosomatidae: Leiobun- inae Phalangium ventricosum ventricosum = Leiobunum ventrico¬ sum ventricosum — — Sclerosomatidae: Leiobun- inae Phalangium verrucosum = Leiobunum verruco¬ sum — — Sclerosomatidae: Leiobun- inae Phalangium vittatum - Leiobunum vittatum — — Sclerosomatidae: Leiobun- inae Platybunus triangularis = Rilaena triangularis — — Phalangiidae: Platybuninae Rilaena triangularis Valid Eastern and western North America Introduced from Europe by 1951 Phalangiidae: Platybun¬ inae, new records from 1 province Strandibunus dorsalis = Mitopus morio — — Phalangiidae: Oligoloph- inae Totals 16 valid North American species in 10 genera and 4 subfamilies; 1 species in Iceland and Europe Canada, northern and central U.S.A., Kalaallit Nunaat, Iceland 10 native, 6 introduced or possibly intro¬ duced from 4 subfamilies 30 new province/ state records, 4 new synonyms, 1 family change, 2 new status, 1 new subgenus Acknowledgments We are grateful to many people for their help. The following curators kindly loaned material from their respective museums: David Barr, Royal Ontario Mu¬ seum (ROM); Gergin A. Blagoev and Allison Brown, Biodiversity Institute of Ontario, University of Guelph; David Blades and Robert Cannings (retired), Royal British Columbia Museum; Jonathan A. Coddington, Ralph E. Crabill, Jr. (deceased), Patricia Gentili-Poole, and Hannah Wood, Smithsonian Institution, United States National Museum (USNM); Rod Crawford (retired), Burke Memorial Washington State Museum (BMM); C. D. Dondale (retired), Canadian National Collection of Insects, Arachnids and Nematodes (CNC); Bert Finnamore and Terry Thormin (retired), Royal Alberta Museum; Saul Frommer (retired), University of California Riverside (UCR); Manfred Grasshoff (retired), Senckenberg Natur-Museum und Forschungsinstitut (SNMF); Jurgen Gruber (retired). 38 Special Publications, Museum of Texas Tech University Naturhistorisches Museum, Wien (NMW); Erling Hauge (deceased), Universitetsmuseet i Bergen; Torb- jorn Kronestedt (retired), Naturhistoriska Riksmuseet, Stockholm (NRS); Herbert Levi (deceased), Museum of Comparative Zoology, Cambridge (MCZ); Andrew Moldenke, Oregon State University (OSU); Gary L. Parsons, Michigan State University, East Lansing (MSU); Norman Platnick (retired), American Museum of Natural History (AMNH); and Geoff Scudder (re¬ tired), Spencer Museum, University of British Colum¬ bia (UBC). GRSM = Great Smoky Mountains National Wildlife Refuge collection at Twin Creeks Science and Education Center. Phil Bragg, Cassie Aitchison (CWA), Dave Blades, Don J. Buckle (DJB), Laurent Lesage, and Seppo Koponen also loaned us specimens from their personal collections or provided other help. The collections recorded in this paper as JCC will be transferred to the AMNH; several of these specimens appear (based on data labels) to have started there in the first place. RGH stands for Robert G. Holmberg as the collector and the personal collection. Gifts of speci¬ mens were appreciated from: Matthew Bowser, Kenai National Wildlife Refuge, Alaska; Maria L. Goodnight (deceased); Daniel T. Jennings (retired), University of Maine at Orono, Maine; Yuri M. Marusik, Institute for Biological Problems of the North, Magadan, Rus¬ sia; and Nobuo Tsurusaki, Tottori University, Japan. Richard Bradley (retired) of The Ohio State University Chelicerates Collection (OSAL) kindly checked for specimens that Mary Walker recorded in 1928. Finally, Gary L. Parsons (MSU) was of considerable help in understanding the specimens and labels of Odiellus housed at that collection. We also thank the follow¬ ing for the use of their photographs to help illustrate our fauna: Matthew Bowser (Kenai National Wildlife Refuge, Alaska), Pierre-Marc Brousseau (Montreal, Quebec), Don J. Buckle (Saskatoon, Saskatchewan), Jan van Duinen (Odoorn, Netherlands), and Joseph V. Higbee (Spanaway, Washington). Finally, we thank the many collectors. Literature Cited Acosta, L. E., A. Perez Gonzalez, and A. L. Tourinho. 2007. Methods for taxonomic study. Pp. 494-505 in The Har¬ vestmen: The Biology of Opiliones (R. Pinto da Rocha, G. Machado, and G. Giribet, eds.). Harvard University Press, Cambridge, Massachusetts. Arthofer, W., H. Rauch, B. Thaler-Knoflach, K. Moder, K. Mus¬ ter, B. C. 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