Reprinted fro m FESTSCHRIFT FOR WALTER W . DALQUEST IN HONOR OF HIs SIXTY-SIXTH BIRTHDAY (N. V. Horner, ed . ) Dept . Biology, Midwestern State Univ ., 1984, xx+163

A NEW GENUS OF NORTH AMERICAN HARVESTME N (ARACHNIDA: : PALPATORES )

JAMES C . COKENDOLPHER

ABSTRACT .Dalquestia, a new genus of harvestmen is described . Eurybunus formosus Banks , from Texas, is redescribed and designated the type species of Dalquestia. Globipes rugosu s Schenkel, from California, is redescribed and transferred to the new genus . Dalquestia concho , n . sp . and Dalquestia grasshoffi, n. sp. are described from Chihuahua—Durango and Hidalgo , Mexico, respectively .

In 1980, Cokendolpher reported that Globipes rugosus Schenkel, Glo- bipes formosus (Banks), and an undescribed species from Mexico differe d from typical Globipes spp. Further studies revealed that these specie s represented an undescribed genus and that a second undescribed specie s exists in Mexico . It is the purpose of this paper to describe the new genus and the two new species . The new genus is a member of an undescribed family or subfamily, her e referred to as the Metopilio assemblage . This group was first noted b y Gruber (1969) and consists of Metopilio Roewer, Diguetinus Roewer , Globipes Banks, Eurybunus Banks, and Dalquestia, new genus . As the classification of the is unresolved, I refrain from formall y describing a new taxon for the assemblage . Current ideas on the classifica- tion of the Phalangioidea embrace essentially the same groups, but th e different systems rank particular taxa at significantly different levels (i .e., a tribe in one system will be a family in another system) . The notion tha t members of the Metopilio assemblage belong to the Leptobunidae (Starega , 1978, and others in correspondence) is incorrect . The Leptobunidae ha s been a catchall group for many years . It is now known that this group i s not monophyletic and that the type genus Leptobunus Banks is a member of the Phalangiinae, (Cokendolpher, 1981b) . It should be noted that Starega (1978) was correct in transferring Globipes charitonov i Gritzenko (from Kazakhskaya S. S. R., central Asia) to Egaenus C . L. Koch . The Metopilio assemblage is apparently restricted to southern Nort h America and Central America .

MATERIALS AND METHOD S Acronyms for collections from which specimens were examined are liste d in the acknowledgments . Specimens in my personal collection are liste d JCC. All anatomical measurements are in millimeters and were obtained using a binocular microscope equipped with an ocular micrometer as outlined b y Cokendolpher (1981a) . Descriptions are based on all available specimens , but measurements were only taken of the largest and smallest (based o n 27

28 FESTSCHRIFT FOR WALTER W . DALQUES T

total length when more than two specimens were available) of each sex . The genitalia were removed from the body, examined and illustrated, and the n placed in genitalia vials, which in turn were placed in the specimen vials . The male genitalia were immersed in 100% glycerol for study . A tuft of cotton placed in glycerol was used to support the penes when lateral view s were desired. The female genitalia were first examined in 100% clove oil an d then in 100% lactophenol (Krantz, 1970 :49). The ovipositors were soaked i n absolute ethyl alcohol (to remove all traces of oil and phenol) prior t o rehydration and placement in genitalia vials . All illustrations were prepare d with the aid of a camera lucida .

Dalquestia, New Genus

Eurybunus : Banks, 1910 :156; Comstock, 1948 :72 (in part) . Not Banks, 1893 :209. Globipes : Roewer, 1912 :270 (in part) ; 1923 :862 (in part) ; 1956 :262 (in part) ; Gertsch, 1939 :20 ; Schenkel, 1951 :55 ; Forcart, 1961 :53 ; Gritzenko, 1972 :1572 (in part) ; Rowland and Reddell , 1976:11 ; Cokendolpher, 1980 :134 (in part). Not Banks, 1893 :208 . Type species.Eurybunus formosus Banks, by the present designation . Etymology.Named in honor of Dr. Walter W. Dalquest ; the gender o f the name is considered feminine. Diagnosis and comparisons .Dalquestia differs from all known genera o f Palpatores by the combination of the following characters : 1) palpal claw s smooth, 2) lateral coxal denticle rows absent, 3) dorsal abdominal scute s fused, with seven rows of tubercles, 4) scent gland pores distinctly visibl e from above, 5) all legs with distinct longitudinal rows of tubercles, 6) tibiae angular in cross section, 7) femora lacking pseudo-articulary nodules, 8 ) penis with base of corpus greatly enlarged, and 9) glans an extension o f corpus, junction immovable . Dalquestia is clearly a member of th e unnamed assemblage in which Gruber (1969) placed the genera Metopilio , Diguetinus, Eurybunus, and Globipes. Like Metopilio and Diguetinus, Dalquestia has rows of abdominal tubercles . Dalquestia differs from Metopilio and Diguetinus primarily by the angular cross section of the le g tibiae, and by having non-alate penes which have the corpus greatl y enlarged. Dalquestia differs from Eurybunus and Globipes (note that there are some misplaced Metopilio spp . described or incorrectly placed i n Eurybunus and Globipes) not only by the presence of abdominal tubercles , but by the characters mentioned above to separate Dalquestia from Metopi- lio and Diguetinus . Although it is of limited use, Dalquestia spp. have one or no pseudosegments on tibiae II. Two or more pseudosegments are know n from other members of the assemblage . Description .Small to medium sized harvestmen with hard granulate bodies; scutes fused, thoracic tergite margins indistinct laterally ; with seven rows of dorsal abdominal tubercles (Figs . 1-5) . Ocular tubercle low, wit h two rows of well developed tubercles . Chelicerae with or without tooth on basal joint ; movable finger without apophysis (Figs . 13 and 14) . Suprachelic- eral lamellae indistinct, not visible from above. Preocular area with a small

COKENDOLPHER—NEW GENUS OF NORTH AMERICAN HARVESTMEN 29

FIG. l .Dalquestia formosa male from cave in central Texas .

30 FESTSCHRIFT FOR WALTER W . DALQUEST

FiGs. 2-5 .-Dalquestia males, dorsal aspects : 2, D . formosa; 3, D . rugosa ; 4, D . grasshoffi ; 5 , D . concho (scale lines = 1 .0 mm) .

COKENDOLPHER—NEW GENUS OF NORTH AMERICAN HARVESTMEN 3 1

8

FiGs . 6-14 .Dalquestia appendages : 6, D . formosa male leg I, lateral aspect ; 7, tibia cros s section ; 8, femur cross section ; 9, D . formosa male leg II, lateral aspect ; 10, tibia cross section ; 11, femur cross section; 12, D . formosa male palpus, mesal aspect ; 13, D . Formosa mal e chelicera, lateral aspect; 14, D . rugosa male chelicera, lateral aspect (scale line = 0 .5 mm fo r figs . 7, 8, 10, 11, and 12 ; 0.58 mm for figs . 13 and 14 ; 1 .0 mm for figs . 6 and 9) .

medium hump, covered with pointed tubercles . Scent gland pores facin g anterodorsad, with paired dark depressions, mounted on small raised areas , and often with paired lateral tubercles . Abdominal sternites closely spaced , borders indistinct . Leg coxae without lateral rows of denticles; I and II I enlarged in males; endites of coxae II extended in front of genita l operculum, forming obtuse angle to each other . Legs short, femora I I approximately equal to (males) or shorter than (females) body length ; psudoarticulary nodules of femora lacking ; tibia II with one or n o pseudosegments; metatarsi without pseudosegments . Leg claws smooth , untoothed . Males with legs I and III enlarged, metatarsi and basal portion s of tarsi with ventral rows of spine-tipped tubercles . Femora and tibia of legs

32 FESTSCHRIFT FOR WALTER W . DALQUEST

(less so on I and II) pentagonal or hexagonal in cross section (Figs . 6-11). Genital operculum lacking lateral rows of denticles . Palpi with many tubercles on all sides except mesal margins, lacking apophyses ; femora, patellae, and tibiae flattened mesally; males with tarsi ventrally armed wit h many small tubercles ; claws untoothed, smooth (Fig. 12). Penes broadest a t base, sharply tapering to a thin shaft ; distal 1 /4 of corpus flattened ; glans rigidly fixed to corpus, with four delicate bristles distodorsally . Stylus thickened and slightly fusiform (Figs . 15-21) . Ovipositors with 25 to 28 rings and three segmented furcaC second furcal segment with two to three sli t sensilla (rarely with single sensillum) per side . Seminal receptacles consist- ing of paired convoluted tubules (Figs . 23-27) . Subordinate taxa.The genus Dalquestia contains four species, D. formosa (Banks), D. rugosa (Schenkel), D. concho, n. sp., and D. grasshoffi , n. sp. In addition to differences in morphology, these species can easily b e distinguished by their distributions, all being widely separated (Fig . 28).

Key to the Species of Adult Dalquestia 1. Tibiae II with distinct pseudosegments; basal segment of chelicerae smooth dorsally 2 Tibiae II lacking pseudosegments ; basal segment of chelicerae with tubercles dorsally (Figs . 13 and 14) 3 2. Tibiae II sharply bicolored; total length of body greater than 4 .30 mm (male) or 5 .60 m m (female) ; Chihuahua and Durango border D . conch o Tibiae II distally lightened but not sharply bicolored ; total length of body less than 3 .50 m m (male) or 5 .20 mm (female) ; Hidalgo D . grasshoff i 3. Tibiae of all legs with distal white bands ; abdominal tubercles approximately uniform i n size (Fig . 2) ; Texas D .formosa Tibiae of all legs uniformly brown or brown mottled with white ; abdominal tubercle s noticeably larger towards the posterior midline (Fig . 3) ; southwestern California . . . .D . rugosa

Dalquestia formosa (Banks), new combinatio n Figs . 1, 2, 6-13, 15, 16, 23, 24, 2 8 Eurybunus formosus Banks, 1910 :156; Comstock, 1948 :72 . Globipes formosus : Roewer, 1912 :270, 1923 :826, 1956 :262 ; Gertsch, 1939 :20 ; Gritzenko , 1972:1574 ; Rowland and Reddell, 1976 :11 ; Cokendolpher, 1980:134 . defTI ;/S Types.—Although Banks (1910) did not give manyit is apparent from examination . of museum specimens that two differen t collections were involved. One collection was made at Barton Creek, nea r Austin, Travis Co., Texas, elev.? (22 Nov . 1899, J. H. Comstock) and was catalogued as Cornell University Lot 255 . I am selecting a male from Lo t 255 as the lectotype . This male and two females, herein designate d paralectotypes, are deposited at the MCZ. The remaining paralectotypes ( 1 male, 1 female, 1 juvenile) forming Lot 255 are part of the Cornel l University collection on semi-permanent loan to the AMNH . The second collection of syntypes, herein designated paralectotypes, are likewise fro m the Cornell University collection on loan to the AMNH and consists of 1

COKENDOLPHER—NEW GENUS OF NORTH AMERICAN HARVESTMEN 3 3

22

2 1

Z1 A B

FiGs . 15-27 .Dalquestia genitalia : 15, D . formosa penis, dorsal aspect ; 16, lateral aspect of distal end ; 17, D. rugosa penis, dorsal aspect; 18, lateral aspect of distal end ; 19, D. grasshoffi penis, dorsal aspect ; 20, lateral aspect of distal end; 21, D . concho penis, dorsal aspect ; 22 , lateral aspect of distal end ; 23, D . formosa seminal receptacle, specimen from Chiso s Mountains ; 24, D . formosa seminal receptacle, specimen from central Texas (Kerr Co.) ; 25, D . rugosa seminal receptacle ; 26, D . grasshof f i seminal receptacle ; 27, D . concho semina l receptacle (scale line A = 0 .1 mm for seminal receptacles, B = 0 .5 mm for penes) .

34 FESTSCHRIFT FOR WALTER W . DALQUES T

male and 2 juveniles collected at Austin, Travis Co ., Texas, 500-600 ft (150 - 180 m) elev. (12-18 March 1903, J . H . Comstock) . Diagnosis and comparisons .In most respects, D. formosa resembles D. concho, n. sp. and D. grasshoffi, n . sp., but can be easily distinguished fro m these two species by its larger size, lack of tibiae II pseudosegments, and by the presence of many tubercles on the leg coxae (only marginal groups of tubercles on the other two species) . Dalquestia formosa and D . rugosa can be separated by the differences in their chelicerae and in color as noted i n the key to species (see also "diagnosis and comparisons" of D. rugosa) . Distribution .Central and southwestern Texas in the U .S.A. (Fig. 28). Description .Males : Body large, total length 5 .67-6.91, greatest widt h 2.72-3.94, maximum height 2 .33-3.10. Dorsum dark brown to black, ofte n appearing metallic, with white to pinkish-white (rarely pink) spots o n median portions of last three abdominal tergites (Fig . 1); rarely with pal e pink to white small dots between first and second abdominal tergites ; lateral margins of cephalothorax (posterior to scent gland pores) and abdomen (anterior to front margin of penultimate tergite) white to yellowish-white ; thoracic and abdominal tubercles large and approximately equal in siz e (Fig. 2). Ocular tubercle concolorous with cephalothorax ; length 0.31-0.46, width 0 .36-0.45, height 0.19-0.20; with 5-7 sturdy, pointed tubercles on eac h side. Venter smooth and creamy-yellow white to light brown ; except for posterior end of abdomen and lateral margins of abdominal sternites , orangish-brown to black (commonly dark brown) with small pointe d tubercles. Genital operculum length 1 .38-2.18, width at base 1 .09-1 .43, width at neck 0.58-0.78. Palpal segment lengths : femora 0 .86-1 .22, patellae 0 .72 1 .12, tibiae 0 .83-1 .39, tarsi 1 .30-1 .74. Chelicerae with distinct ventral spur o n basal segment, with few tubercles dorsally (Fig . 13) . Chelicerae and palpi light brown to black. Legs (Figs. 6-11) with distinct rows of large pointe d tubercles on femora, patellae, and tibiae ; tibiae II without pseudosegments . Legs yellowish-white to light yellowish-brown with distal parts of femora I I shaded yellowish-brown to brown ; distodorsal tips of femora II and IV an d all tibiae white ; distal parts of patellae shaded brown ; all tibiae, metatars i II, and tarsi II and IV dark brown . Tarsi I and III, metatarsi I, III, and I V dark yellowish-brown ; distal tips of tarsi and metatarsi white ; proximal end s of metatarsi IV, rarely III, white . On occasion leg femora (excluding II), al l coxae, and trochanters with considerable irregular white reticulations . Coxae with lateral clusters of tubercles on anterior and posterior margins of I, III, IV, and anterior of II ; few scattered small tubercles on all coxae , particularly numerous on I . Femora I-IV lengths (respectively) : 2.28-2.63 , 5.68-6.15, 2.18-2.67, 3 .32-3.70; tibiae I-IV lengths (respectively): 1.97-2.58 , 4.61-5.14, 1 .82-2.38, 1 .82-2.82. Penis as in figures 15 and 16, length 2 .72-2.86. Females : Form and coloration as in males except palpi and legs no t enlarged or modified, tubercles reduced in size and number on leg coxa e and chelicerae dorsally . Lateral white stripes of dorsum extend from jus t posterior of scent gland pore to anterior edge of penultimate abdominal

COKENDOLPHER—NEW GENUS OF NORTH AMERICAN HARVESTMEN 35

FIG . 28 .—Distribution of Dalquestia, star = D . rugosa, circle = D. formosa, triangle = D . concho, square = D . grasshof f i .

tergite. Total length 4 .58-7.16, greatest width 3 .12-4.19, maximum heigh t 2.16-3.00. Ocular tubercle longer than wide, length 0 .38-0.39, width 0 .32- 0.35, height 0.14-0.17, with five large rounded tubercles over each eye . Genital operculum length 1 .40-1 .48, width at base 1 .34-1 .68, width at neck 0.74-0.76. Palpal segment lengths : femora 0.78-0.80, patellae 0 .55-0.57, tibia e 0.64-0.66, tarsi 1 .15-1 .33. Tibiae II without pseudosegments . Femora I-I V lengths (respectively) : 2.00-2.08, 5 .61-6.08, 2.06-2.08, 3.43-3 .75; tibiae I-IV lengths (respectively) : 1.58-1 .78, 4.31-4.54, 1 .74-1 .82, 2.31-2.49. Seminal receptacles as in Figures 23 and 24. Immatures : Body smooth, lengths 3 .3-4 .5 ; dorsally reddish-brown with dark reddish-brown ocular tubercle ; creamy-white stripes laterally from scen t gland pores to about middle of abdomen and creamy-white spot medially on last three abdominal tergites ; with characteristic rows of tubercles, as in

36 FESTSCHRIFT FOR WALTER W . DALQUEST

adults . Venter creamy-white to whitish-yellow except for anal plate and lateral portions of last three to four sternites reddish-brown . Chelicerae and palpi creamy-white with extensive reddish-brown mottlings . Basal joint of chelicerae with distinct ventral spur. Palpi without apophyses or tubercles . Leg coxae, trochanters, and bases of all femora creamy-white ; femora (excluding II) and patellae (excluding II) light orange to creamy-orang e distally, femora II and patellae II distally orangish-brown . Extreme dista l tips of femora II orange, all tibiae reddish-brown with broad white band s on II and creamy-white to whitish-orange on distal ends of I, III, and IV . Tarsi and metatarsi light creamy-brown to light reddish-brown with whit e on proximal ends of metatarsi III, and distal ends of metatarsi III and IV . Natural history.From labels accompanying museum specimens, it i s clear that many of the specimens were collected under rocks and pieces o f wood . Adult specimens have been collected throughout the year with th e majority being taken from July to November . Juveniles are likewise known throughout the year, but too few specimens are known to determine an y seasonal trends. Two females collected during October contain large eggs . This species is known from essentially two regions of Texas, the Chiso s Mountains and central Texas. From the latter region, specimens have bee n collected at altitudes from 150 to 550 m, with the majority from 290 to 36 0 m elevation. Collections from the Chisos Mountains were made at elevation s from 1370 to 2130 m, with the majority from 1550 to 1640 m . No othe r natural history data are available . Specimens examined.U .S .A ., TExAS : county ?, 700 Springs (2 April 1936, collector ?), 1 female (AMNH) ; Palo Pinto Co ., Possum Kingdom State Fish Hatchery (21 Oct . 1979, E . Matelski), 2 males, 1 female (JCC), 3 mi (4 .8 km) W . of S.H . 16 on P36, Possum Kingdom Lake (21 Oct . 1978, G . and C . Spicer), 1 male, 1 female (JCC), 250 yds (288 m) W . of Brazos River on Hwy . 180 (8 Dec . 1973, W . R . Barber et al .), 1 female, 1 juvenile (JCC) ; San Saba Co. , specific locality ? (5 may 1973, T . C. Kaspar), 1 female (JCC) ; Travis Co., Barton Creek (22 Nov . 1899, J . H . Comstock), lectotype male, 2 female paralectotypes (MCZ-Cornell Lot 255), 1 male, 1 female, 1 juvenile paralectotypes (AMNH-Cornell Lot 255), Austin (12-18 March 1903 , J . H . Comstock), 1 male, 2 juvenile paralectotypes (AMNH), Travis Park, Lake Travis (1 0 March 1946, H . E . Frizzell), 1 male, 1 female (SMF-RII/9050/301) ; Comal Co., specific locality ? (16 Oct . 1951, D. L . Frizzell, H . E . Frizzell, and S . B . Martin), 1 male (EP), U .S . Hwy . 281 nr. Guadalupe River (24 Sept . 1939, Bogert and Vokes), 1 male, 1 penultimate female (AMNH) ; Bexar Co ., 10 mi (16 km) N . of San Antonio (18 July 1942, E . S. Ross), 2 males, 1 femal e (CAS), (31 Jan . 1943, E . S . Ross), 1 male, 3 females (CAS), 20 mi (32 km) N . of San Antoni o (13 Dec . 1942, E . S . Ross), 3 males, 1 female, 1 juvenile (CAS) ; Kendall Co ., Century Caverns (Cave Without-A-Name) (14 Aug. 1966, K . Hom and A . Jung), 2 males, 1 female (CAS) ; Ker r Co ., specific locality ? (27 Oct . 1979, N . V . Horner), 1 male (MSU), Camp Chrysalis, 14 m i (22.5 km) S . W . of Kerrville (15 Oct . 1977, D . Holub and J . C . Cokendolpher), 1 male, 1 female , (MSU), 5 males, 5 females, 2 juveniles (JCC) ; Edwards Co . 15 mi (24 km) W . of Rock Spring s (22 March 1979, O. F . Francke, J . V . Moody, W . D . Sissom, and F . Merickel), 1 penultimate female (OFF); Kimbell Co ., 7 mi (11 km) E . of Junction (21 Sept. 1968, R . Carpenter), 1 mal e (MSU) ; Schleicher Co ., 3 mi (4 .8 km) N .W . of Eldorado (10 Aug . 1978, O. F. Francke, J . V . Moody, and B . Hall), 1 female, 1 juvenile (OFF) ; Brewster Co ., Big Bend National Park, Pin e Canyon (10 Aug. 1979, O . F . Francke, J . V . Moody, and F . Merickel), 1 female (OFF) ; Kibbee Spring (11 April 1945, K. P . Schmidt), 1 male (FMNH), Juniper Canyon (5 Aug . 1937, K . P . Schmidt), 1 juvenile (FMNH), CCC Camp, Chisos Basin (25-30 July 1937, W . L . Necker), 1

COKENDOLPHER—NEW GENUS OF NORTH AMERICAN HARVESTMEN 3 7

male, 1 female (FMNH), Chisos Basin (2 Aug . 1938, collector ?), 3 males (AMNH), 3 male s (ALE), (31 July 1937, K . P. Schmidt), 1 male, 1 female (FMNH), (6 Aug . 1937, K. P . Schmidt) , 1 juvenile (FMNH), Chisos Basin Pass (28-29 July 1979, O . F . Francke and J . V . Moody), 1 male, 1 penultimate male, 2 females, 2 juveniles (OFF) .

Dalquestia rugosa (Schenkel), new combination Figs . 3, 14, 17, 18, 25, 28 Globipes rugosus Schenkel, 1951 :55, fig . 49; Forcart, 1961 :53 ; Cokendolpher, 1980 :134 . Types.Male lectotype and male (reported as a juvenile) paralectotype (designated by Forcart, 1961 :53) from Country Club Heights, La Jolla, Sa n Diego Co ., California, 500 ft (150 m) elev. (10 Jan. 1939, H. Schenkel - Rudin), NMB no. 90-a, examined . Diagnosis and comparisons .In addition to the characters stated in th e key to species, D. rugosa can be distinguished by the lack of white spots o n the posterior end of the abdomen, which are so striking in other Dalquesti a spp. (Fig. 1). The cheliceral spur or "hook" is not developed in D. rugosa as it is in its congeners (Figs . 13 and 14), but the development of tubercles o n the distal segment of the chelicerae occurs only in D. rugosa males. Distribution .Known only from southwestern California in the U .S.A. (Fig. 28). Description .Males : Body large, total length 4 .67-5.81, greatest widt h 2.42-3.53, maximum height 2 .38-2.64. Dorsum roughly granulated, ligh t brown to dark reddish-brown with considerable spotting and mottling o f darker brown ; small tubercles and ridges of abdomen reflect light a s sparkles; rounded tubercles on abdomen increase in size posteriorly an d medially (Fig . 3). Ocular tubercle length 0 .38-0.41, width 0 .35-0.42, heigh t 0.12-0.18, with 4-7 short rounded tubercles on each side . Venter uniforml y light yellow brown to light brown . Anal plate and posterior half of the las t two sternites strongly rugose ; anterolateral margins of all abdomina l sternites rugose . Genital operculum length 1 .46-2.07, width at base 1 .03 - 1 .58, width at neck 0 .43-0.71 . Palpal segment lengths : femora 0 .79-1 .12, patellae 0 .60-0.78, tibiae 0 .72-0.98, tarsi 1 .19-1 .47. Chelicerae with man y tubercles on dorsal surfaces of both segments ; basal joint without ventral spur (Fig . 14) . Legs, palpi, and chelicerae light yellowish-brown to ligh t reddish-brown in color. Leg femora and tibiae slightly darker in color an d mottled, with rows of large tubercles . Coxae evenly covered with pointed tubercles except anterior edges of III slightly more tuberculate . Tubercles o n leg coxae, trochanters, palpi, and chelicerae dark brown to black, ver y conspicuous . Femora I-IV lengths (respectively) : 1 .80-2.50, 4.28-5 .80, 1 .62- 2.09, 2.16-3.08; tibiae I-IV lengths (respectively) : 1 .69-2.48, 3.81-5 .00, 1 .95- 2.00, 1 .83-2.38. Penis as in figures 17 and 18, length 3 .18-4.02. Females : Form and coloration as in males, except legs, palpi, and chelicerae not enlarged or modified, and body not as tuberculate . Chelicerae with few tubercles dorsally on basal joint, none on distal joint, lackin g distinct spur ventrally . Tubercles on abdomen dorsum much smaller than

38 FESTSCHRIFT FOR WALTER W . DALQUEST

on males (largest female tubercle slightly larger than smallest tubercle o n males). Total length 5.28-5 .49, greatest width 2 .71-2.80, maximum heigh t 1 .76-2.18. Ocular tubercle slightly longer than wide, length 0 .27-0.34, width 0.25-0.30, height 0.13-0.14, with 5-7 low tubercles on each side . Genita l operculum length 1 .10-1 .15, width at base 1 .14-1 .26, width at neck 0.37-0.56. Palpal segment lengths : femora 0.66-0.70, patellae 0.35-0.43, tibiae 0 .50-0.56. tarsi 0.93-1 .06. Tibiae II without pseudosegments . Femora I-IV length s (respectively) : 1 .60-1 .72, 4.50-?, 1 .42-1 .89, 2.63-2.81 ; tibiae I-IV length s (respectively) : 1 .31-1 .40, 3 .77-?, 1 .26-1 .31, 1 .63-1 .67. Seminal receptacle as i n figure 25 . Immature: Body length 4 .4; dorsally uniform light brown with dar k brown to black rings around eyes ; smooth except for two thoracic and seve n abdominal rows of tubercles . Venter smooth except for small tubercles o n anal plate ; light creamy-yellow . Palpi and chelicerae lacking all tubercles , light yellowish-brown to light reddish-brown in color . Basal segment o f chelicera with rounded ridge, spur absent . Legs with tibiae angular in cross section ; with rows of small tubercles on femora, patellae, and tibiae ; tibiae II without pseudosegments ; light yellow brown to reddish-brown with som e mottling . Natural history .All known specimens, except for the types, wer e collected with pitfall traps . Adults have been taken throughout the year . Two females collected on 15 January and 29 August, respectively, had thei r abdomens distended and contained large eggs . The single juvenile, a late instar female, was collected on 19 December . Collections have been made in two different regions about 100 km apart . Near the type locality, specimen s come from areas between 100 and 150 m in elevation and are found i n altered (disturbed) chaparral . Other samples are known from Paloma r Mountain (550-1350 m elevation). Specimens examined .U .S .A., CALIFORNIA : San Diego Co., Country Club Heights, La Joll a (10 Jan . 1939, H . Schenkel-Rudin), male lectotype, male paralectotype (NMB no . 90-a) ; Sa n Diego, Hewlett Drive (Oct . 1970, B . J . Kaston), 1 male (WAS) ; Palomar Mountain, Nat e Harrison Grade, Banta Station 45 (18 April 1970, B . H . Banta), 1 male (CAS), (17 July 1971, B . H . Banta), 1 male (CAS), (15 Jan . 1972, B . H . Banta), 1 female (CAS) ; Banta Station 52 (2 9 Aug . 1970, B . H . Banta), 1 female (CAS) ; mixed stations (2063-4481 ft elev .—19 Dec . 1970, B . H . Banta), 1 juvenile female (CAS) .

Dalquestia grasshoffi, new specie s Figs . 4, 19, 20, 26, 2 8 Types.—Male holotype, male paratype, and two female paratypes fro m Jacala, Hidalgo, Mexico, 4000 ft (1200 m) elev. (20 April 1963, W . J. Gertsch and W . Ivie), AMNH . Etymology .The specific epithet is a patronym in honor of Dr . Manfred Grasshoff for his contributions to arachnology . Diagnosis and comparisons .Dalquestia grasshoffi is most similar to D. concho, and from that species, it can be separated by its smaller size and

COKENDOLPHER—NEW GENUS OF NORTH AMERICAN HARVESTMEN 3 9

differences in the leg tibiae . The femora and tibiae of D. grasshoffi are concolorous, except for the distal ends of the tibiae which are lighter i n color (not distinctly bicolored) . The tubercles on tibiae II are reduced in siz e on D. grasshoffi . The basal portions of the leg tibiae in D. concho are always darker in color than the femora . Tibiae II always have distinctiv e white bands distally . For additional comments, see diagnosis of D. formosa. Distribution .Known only from Jacala, Hidalgo, Mexico (Fig. 28). Description .Males : Body small, total length 3 .09-3.45, greatest width 2.00-2 .18, maximum height 1 .34. Dorsum light reddish-brown with smal l white opalescent spots on middle of abdominal tergites I and II and large r white to silvery-white spots on last three tergites ; thoracic and abdomina l tubercles rounded and approximately equal in size (Fig . 4). Ocular tubercl e dark reddish-brown in color ; longer than wide, length 0 .27-0.30, width 0.23- 0.25, height 0 .12-0.18; with 5-6 tubercles on each side . Genital operculu m and venter essentially smooth, except for small pointed tubercles on latera l margins of abdomen, last free abdominal sternite, and anal plate . Genita l operculum length 0 .95-0.96, width at base 0 .98-1 .00, width at neck 0.43-0.50. Palpal segment lengths : femora 0.72-0.96, patellae 0 .48-0.50, tibiae 0 .60-0.61 , tarsi 0.84-0.96. Chelicerae with distinct ventral spur on basal segment , lacking dorsal tubercles . Venter of body and chelicerae light yellowish- brown ; except anal plate slightly darker, concolorous with dorsum o f abdomen . Palpi and legs yellowish-brown to reddish-brown : coxae slightly lighter, more yellow in color ; distal tips of leg tibiae dorsally with yellowish-white spots (sometimes weak or absent on II) . Leg coxae with fe w pointed tubercles clustered on anterior and posterior margins of I, III, an d IV; II sometimes with one or two tubercles . Tibiae II with rows of smal l tubercles on ventral side and smaller and more greatly spaced tubercles o n basal 1 /3 dorsolaterally ; all other leg femora and tibiae with complete rows of large pointed tubercles. Tibia II with single pseudosegment . Femora I-I V lengths (respectively) : 1 .60-1 .70, 4.81-5.24, 1 .38-1 .84, 2.34-2.44; tibiae I-IV lengths (respectively) : 1 .38-1 .52, 3.82-4.04, 1 .27-1 .29, 1 .69-1 .76. Penis as in figures 19 and 20, length 1 .92-2.04. Females : Form and coloration as in males, except abdominal tergite I I without white spot, femora IV basally yellowish-white, palpal tarsi withou t ventral rows of tubercles, and legs I, III, and palpi not enlarged . Tota l length 4 .00-5.07, greatest width 2 .57-2.80, maximum height 1 .61-1 .93 . Ocular tubercle longer than wide, length 0.32-0.34, width 0.27, height 0.12- 0.20, with 6-8 tubercles on each side . Genital operculum length 1 .04-1 .07 , width at base 1 .12, width at neck 0.52-0.53. Palpal segment lengths : femora 0.61-0.63, patellae 0.36-0.40, tibiae 0 .50-0.52, tarsi 0 .92-0.96. Tibia II with single pseudosegment . Femora I-IV lengths (respectively) : 1 .52-1 .55, 5.06- 5.21, 1 .50-1 .55, 2.66-2.70; tibiae I-IV lengths (respectively) : 1 .29-1 .40, 4.05- 4.73, 1 .26-1 .29, 1.84-1 .91 . Seminal receptacle as in figure 26 . Immatures : unknown .

40 FESTSCHRIFT FOR WALTER W . DALQUES T

Natural history.The only known specimens were collected as adult s during the middle of April at an elevation of 1200 m. The larger of the tw o females contains large eggs . Specimens examined .Male holotype, male paratype, and two female paratypes .

Dalquestia concho, new specie s Figs . 5, 21, 22, 27, 2 8 Globipes n . sp ., Cokendolpher, 1980 :134 . Types.—Male holotype and two female paratypes from 1 .6 km E. of th e ranch LaSauceda, Chihuahua, Mexico, 7000 ft (2100 m) elev . (21 July 1947 , W. J . Gertsch), AMNH ; and eight additional paratypes listed unde r specimens examined. The ranch La Sauceda is located approximately 34 k m (by road) NW of Santa Barbara in a narrow valley surrounded b y mountains . The type locality is near the crest of the mountains (see Spieth , 1950:26, fig. 52). Etymology.The specific epithet is a noun in apposition ; the name of the Indians of the region. Diagnosis and comparisons.In addition to characters stated in the key to species, see diagnosis under D. grasshof f i. Distribution .Southern Chihuahua and northern Durango in Mexic o (Fig. 28). Description .Males : Body small, total length 4.42-4.76, greatest width 2.58-2.78, maximum height 2 .00-2.18. Dorsum reddish-brown to dark brow n with yellow-white longitudinal stripes from just posterior to scent glan d pores to tip of abdomen laterally; last abdominal tergite with a centra l white spot ; thoracic and abdominal tubercles low, rounded, and approxi- mately equal in size (Fig. 5). Ocular tubercle reddish-brown in color ; longer than wide, length 0 .36-0.40, width 0 .32-0.33, height 0 .16-0.19; with 4-6 lo w rounded tubercles on each side . Genital operculum and venter essentiall y smooth, except for small pointed tubercles on anal plate, posterior half o f last abdominal sternite, and sometimes lateral edges of anterior sternites . Genital operculum length 1 .25-1 .36, width at base 1 .37-1 .47, width at neck 0.56-0.58. Palpal segment lengths : femora 0.92-1 .06, patellae 0.68-0.81, tibiae o.89-1 .09, tarsi 1 .17-1 .18. Chelicerae with distinct spur on basal segmen t ventrally, lacking dorsal tubercles . Venter of body, chelicerae, and palpi light yellowish-brown ; except for anal plate darker; palpal femora, patellae , and tibiae dorsally with white reticulations . Legs light yellow to yellowish - brown with distodorsal tips of all leg femora and often patellae with white splotches ; all tibiae, patellae II concolorous with adbominal dorsu m (reddish-brown to dark brown) ; distal tips of tibiae I, III, and IV, and all o f tibiae II distal to pseudosegments white ; metatarsi II dark brown; rows o f pointed tubercles well developed on all femora and tibiae . Coxae of leg s with clusters of pointed tubercles on distal ends of I and IV (anterior an d posterior margins) and III (anterior only) ; other margins with only 2- 5 scattered tubercles . Tibia II with single pseudosegment . Femora I-IV lengths

COKENDOLPHER—NEW GENUS OF NORTH AMERICAN HARVESTMEN 4 1

(respectively): 2.09-2.30, 4.84-5.35, 1 .92-2.74, 2.73-2.96; tibiae I-IV lengths (respectively): 1 .92-2.00, 4.22-4.58, 1 .64-1 .80, 2.00-2.18. Penis as in figures 2 1 and 22, length 2 .37-2.78. Females : Form and coloration as in males, except legs I, III, and palp i not enlarged . White spots on abdominal tergites I and II not well developed . Tubercles on abdomen smaller than on males . Coxae with clusters o f denticles on both sides I, and anterior III and IV; other surfaces eithe r smooth or with only few scattered tubercles . Tubercles distal to pseudoseg- ments on tibiae II (dorsal and dorsolateral surfaces) smaller and mor e greatly spaced than proximal tibiae II tubercles . Total length 5 .71-5.84, greatest width 3 .17-3.22, maximum height 2 .30-2.32. Ocular tubercle longe r than wide, length 0 .32-0.32, width 0.25-0.29, height 0 .15-0.15, with 5-6 large tubercles over each eye . Genital operculum length 1 .13-1 .23, width at bas e 1.18-1 .32, width at neck 0.56-0.61 . Palpal segment lengths : femora 0 .63-0.69, patellae 0 .35-0.44, tibiae 0 .55-0.56, tarsi 0 .97. Tibia II with single pseudoseg- ment . Femora I-IV lengths (respectively) : 1 .67-1.84, 4.96-5 .09, 1 .75-1 .76, 2.82; tibiae I-IV lengths (respectively) : 1 .50-1 .52, 3.90-4.11, 1 .38-1 .48, 1 .80-1 .92. Seminal receptacle as in figure 27 . Immatures : Body lengths 2 .3 and 2.8; dorsally smooth except for rows of large pointed tubercles on thoracic tergites and abdomen (as in adults) an d small to minute tubercles scattered over entire dorsum . Venter, coxae , chelicerae, and palpi smooth. Palpi with few small spines. Chelicerae wit h distinct pointed ventral spur on basal joint . Legs essentially as in adult females but with tubercles greatly reduced in size ; tibia II with singl e pseudosegment . Coloration of body and legs as in adults except much paler ; dorsum yellow brown to light brown . Natural history.—Adults are known from January and July . One of the two females contained large eggs . Although only one specimen wa s collected at some time other than July, it must be remembered that all the July collections were made by a single expedition . Most were collected in mountainous areas with grasses, century plants, a few shru b junipers, and some cacti . Other specimens were taken in moister situation s in sparse oak and pine woods . Photographs of the region as well a s descriptions of each locality are given by Spieth (1950:25-33, figs . 15, 18) . All collections were at elevations from about 1700 to 2300 m, with th e majority from about 1950 m .

Specimens examined .Holotype and 10 paratypes—MEXICO, CHIHUAHUA : 1 .6 km E . o f ranch La Sauceda (21 July 1947, W. J . Gertsch), male holotype, 2 females (AMNH) ; Valle d e Olivos (20 July 1947, W . J . Gertsch), 1 male (AMNH) ; Santa Barbara (20 Jan . 1947, M. G . Bradt), 1 male (AMNH) . DURANGO : Encino (27 July 1947, W . J . Gertsch), 1 male, 1 juvenil e (AMNH) ; Los Puentes (23 July 1947, W . J. Gertsch), 3 males, 1 juvenile (AMNH) .

ACKNOWLEDGMENTS

I am grateful to Drs. Arthur F. Beyer and Norman V . Horner for giving me the opportunity to join former students in honoring Dr. Walter W . Dalquest. Dr . Robert W . Mitchel l generously provided the photograph of Dalquestia formosa, and Dr . Willis J . Gertsch kindl y supplied me with information on his collections from Mexico, including a loan of the

.12 FESTSCHRIFT FOR WALTER W . DALQUEST

publication by Spieth . Thanks are extended to my wife, Jean, for her assistance in all phases o f this study and to Dr . B . J . Kaston for information on the natural history of Dalquestia rugosa . Dr. Oscar F. Francke and Mr . Greg S . Spicer are gratefully acknowledged for their critica l reviews of the manuscript and for gifts and loans of specimens . Thanks are also extended to th e following individuals and curators for their continued assistance with loans of specimens : Dr . Arlan Edgar, Alma College, Alma, Michigan (ALE) ; Dr . Oscar F . Francke, Texas Tec h University (OFF) ; Dr. Manfred Grasshoff, Natur-Museum Senckenberg-Forschungsinstitut , Frankfurt, West Germany (SMF) ; Dr. Norman V . Horner, Midwestern State University (MSU) ; Dr. David H . Kavanaugh, Dr . Wojciech J . Pulawski, and Mr . Vincent F . Lee, Californi a Academy of Sciences, San Francisco, California (CAS) ; Dr . Herbert W . Levi, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts (MCZ) ; Dr . William B . Peck, Exline-Peck Collection, Warrensburg, Missouri (EP) ; Dr . Norman I . Platnick, America n Museum of Natural History, New York, New York (AMNH) ; Dr. William A . Shear, Hampden - Sydney College, Hampden-Sydney, Virginia (WAS) ; Dr . Eric H . Smith, Field Museum of Natural History, Chicago, Illinois (FMNH) ; and Dr . Ernst Sutter, Naturhistorisches Museum , Basel, Switzerland (NMB) . This study was supported in part by funds provided by th e Biology Program at Texas Tech University .

LITERATURE CITE D

BANKS, N . 1893 . The Phalanginae of the United States . Canadian Entomol ., 25 :205-211 . . 1910 . A Handsome Species of Phalangidae . Entomol . News, 21 :156 . COKENDOLPHER, J . C . 1980 . Comments on Opiliones described from Western North Americ a by Schenkel . Entomol . News, 91 :133-135 . 1981a . Revision of the genus Trachyrhinus Weed (Opiliones, Phalangioidea) . J . Arachnol ., 9 :1-18 . 1981b . The harvestman genus Liopilio Schenkel (Opiliones : Phalangiidae) . J . Arach- nol ., 9 :309-316 . COMSTOCK, J . H . 1948 . The spider book . Rev. ed . (W . J . Gertsch, ed .) . Comstock Publish- ing Co ., Inc., Ithaca, 729 pp . FORCART, L . 1961 . Katalog der Typusexemplare in der Arachnida-Sammlung des Naturhistor- ischen Museums zu Basel : Scorpionidea, Pseudoscorpionidea, Solifuga, Opilionidea , and Araneida . Verhandl . Naturf. Ges . Basel, 72 :47-87 . GERTSCH, W . J . 1939. Report on a collection of Arachnida from the Chisos Mountains . Con- trib . Baylor Univ . Mus ., 24 :17-26 . GRITZENKO, N . I . 1972 . Globipes charitonovi sp . n . (Opiliones, Phalangiinae) from Kazakh- stan . Zool . Zhur ., 51 :1572-1574 . GRUBER, J . 1969 . Bemerkungen zur Genitalmorphologie and systematischen Stellung vo n Metopilio australis (Banks) (Phalangiidae : Opiliones, Arachnida) . Ann . Naturhist . Mus. Wien, 73 :271-274 . KRANTZ, G . W . 1970 . A Manual of Acarology . Oregon State Univ . Book Stores, Inc. , Corvallis, 335 pp. ROEWER, C. F . 1912 . Revision der Opiliones Palpatores ( = Opiliones Plagiostethi) II . Teil : Familie der Phalangiidae . (Subfamilien : Sclerosomini, Oligolophini, Phalan- giini) . Abhandl . Naturw. Ver ., Hamburg, 20 :1-295 + 4 pls . . 1923 . Die Weberknechte der Erde . Systematische Bearbeitung der bisher bekannte n Opiliones . Gustav Fischer, Jena, 1116 pp . 1956 . Uber Phalangiinae (Phalangiidae, Opiliones Palpatores) . (Weitere Weber- knechte XIX) . Senckenberg. Biol ., 37 :247-318 . ROWI .AND, J . M ., AND J . R . REDDELL . 1976 . Annotated checklist of the fauna o f Texas (excluding Acarida and Araneida) . Occas . Papers Mus ., Texas Tech Univ . , 38 :1-25 .

COKENDOLPHER—NEW GENUS OF NORTH AMERICAN HARVESTMEN 4 3

SCHENKEL, E . 1951 . Spinnentiere aus dem westlichen Nordamerika, gesammelt von Dr . Han s Schenkel-Rudin . Verhandl . Naturf. Ges . Basel, 62 :28-62 . SPIETH, H . T . 1950 . The David Rockefeller Mexican Expedition of the American Museum o f Natural History . Introductory Account . Amer. Mus . Novitates, 1454 :1-67 . STAREGA, W . 1978 . Katalog der Weberknechte (Opiliones) der Sowjet-Union . Fragm . Faun . , Warszawa, 23 :197-234.

Address of author : Department of Biological Sciences, Texas Tech University, Lubbock , Texas 79409.