CLASSIFICATION, PHTLDSOilY A'MD ZDDffiDSRAPHY Df T-HE

AMERICAM ARAOilllOS OF THE DIDO SCfllZDMIDA

!by

JON WM. KDWLATO, B.S.

A iSISSERTATIUN

ITkl

Subiritted to tte firadiats Faculty of Texas Tecti UniveTsi'ty in Partiiail Fiuilfillment ©f tte Iteqiuiiiirenffiimts for tltne !Die§ree ©f

Accepted

©ecenlber, 1575 ACKNOWLEDGMENTS

Great appreciation is expressed to Dr. Robert W. Mitchell who, by his consistant example of excellence, has constructed and

shaped me as a zoologist, and thus shall share credit for production of this dissertation. The remainder of my committee afforded me

generously of their time and advice and are here thanked: Dr. Russell

W. Strandtmann, Dr. John S. Mecham, Dr. John E. George, and Dr. David

E. Foster. Additional advice was provided by Dr. William R. Atchley.

I am further indebted to the Society of Sigma Xi for awarding this

study a Grand-in-Aid of Research. Fellow students who have

contributed significantly to my dissertation are William R. Elliott,

Jerry W. Cooke, and D. Craig Rudolf. Mr. James R. Reddell has given

freely of his advice and time, and has in all areas contributed his

remarkable knowledge of faunistic zoology in general and schizomids

in particular. Dan, Verlee, and Phyllis Brashier Rowland saw this

venture to completion.

n TABLE OF CONTENTS Page ACKNOWLEDGMENTS ii LIST OF TABLES iv LIST OF FIGURES vi LIST OF MAPS ix I. INTRODUCTION 1 II. MATERIALS AND METHODS 16 III. CLASSIFICATION 22 IV. PHYLOGENETICS 315 V. ZOOGEOGRAPHY 374 BIBLIOGRAPHY 406

111 LIST OF TABLES

TABLE Page

1. Comparisons of the species groups of the American members of the genus Schizomus , 39

2. Measurements of the members of the Protoschizomidae 49

3. Comparisons of the members of the cubanicus group . 52

4. Measurements of the members of the cubanicus group . 95

5. Comparisons of the members of the simonis group . . 101

6. Measurements of the members of the simonis group . . 125

7. Comparisons of the members of the brasiliensis group 130

8. Measurements of the members of the brasiliensis group 161

9. Comparisons of the members of the mexicanus group 166

10. Descriptive statistics for S^. portoricensis . . . 205

11. Measurements of the members of the mexicanus group 229

12. Comparisons of the members of the pecki group . . 234

13. Measurements of the members of the pecki group . . 253

14. Comparisons of the members of the goodnightorum group 257

15. Measurements of the members of the goodnightorum group 268

16. Comparisons of the members of the briggsi group . . . 271

17. Measurements of the members of the briggsi group . . . 297

18. Measurements of the species of Schizomus not assigned to groups 313

19. Characters, character states and assigned codes of the major lineages of Schizomida 342

20. Data matrix of the genera 344

21. Characters, character states and assigned codes of the non-briggsi American species groups 347

TV TABLE Page

22. Data matrix of the non-briggsi American species groups . . 348

23. Characters, character states and assigned codes of the members of the cubanicus group 351

24. Data matrix of the members of the cubanicus group .... 353

25. Characters, character states and assigned codes of the members of the simonis group 356

26. Data matrix of the members of the simonis group 357

27. Characters, character states and assigned codes of the members of the brasiliensis group 360

28. Data matrix of the members of the brasiliensis group . . . 362

29. Characters, character states and assigned codes of the members of the mexicanus group 365

30. Data matrix of the members of the mexicanus group .... 367

31. Characters, character states and assigned codes of the members of the briggsi group 370

32. Data matrix of the members of the briggsi group 371 LIST OF FIGURES

FIGURE Page

1. Male and female Agastoschizomus lucifer from Sotano de Yerbaniz, San Luis Potosi 9 2. Male and female Schizomus pentapeltis from near Palm Springs, California, during courtship. Male S^. mexicanus from Sotano de la Tinaja, San Luis Potosi . 11

3-6. Generalized schizomidan anatomy 13

7-13. Generalized schizomidan anatomy 15

14-19. Parts of protoschizomids 47

20-28. Dorsal views of male's flagella of the cubanicus group 79 29-33. Dorsal views of male's flagella of S^. viridis from various localities 81

34-44. Male's flagella of the cubanicus group 83

45-57. Parts of male schizomids of the cubanicus group ... 85

58-60. Lateral views of male's right pedipalps of the cubanicus group 87 61-68. Lateral views of male's right pedipalp of the cubanicus group 89

69-73. Male's right pedipalps of the cubanicus group .... 91

74-86. Female's spermathecae of the cubanicus group .... 93

87-94. Dorsal views of male's flagella of the simonis

group 119

95-106. Parts of schizomids of the simonis group 121

107-118. Female's spermathecae of the simonis group 123

119-125. Dorsal views of male's flagella of the brasiliensis

group 153 126-136. Male's flagella of the brasiliensis group 155

137-145. Parts of schizomids of thVI e brasiliensis group .... 157 FIGURE Page

146-154. Female's spermathecae of the brasiliensis group . . . 159

155-163. Dorsal views of male's flagella of the mexicanus

group 215

164-170. Parts of schizomids of the mexicanus group 217

171-182. Lateral views of male's flagella of the mexicanus

group 219

183-190. Female's spermathecae of the mexicanus group .... 221 191-198. Female's spermathecae of the mexicanus group .... 223

199-211. Parts of schizomids of the mexicanus group 225

212-215. Male's pedipalps of S.. mitchelli 227

216-221. Parts of schizomids of the pecki group 247

222-228. Female's spermathecae of the pecki group 249

229-234. Parts of schizomids of the pecki group 251

235-240. Parts of male schizomids of the goodnightorum group . 264

241-247. Parts of schizomids of the goodnightorum group . . . 266

248-253. Male's flagella of the briggsi group 289

254-259. Parts of schizomids of the briggsi group 291

260-265. Female's spermathecae of the briggsi group 293

266-272. Parts of male schizomids of the briggsi group .... 295

273-279. Parts of male schizomids 309

280-284. Parts of schizomids 311

285. Schematic diagram of the probable cladistic

relationships of the flagella of schizomidans .... 330

286-287. Cladograms of the genera of schizomidans 346

288. Cladogram of the non-briggsi groups of American schizomids vn 350 FIGURE Page

289. Cladogram of the members of the cubanicus group 355

290. Cladogram of the members of the simonis group 359

291. Cladogram of the members of the brasiliensis group . . . 364

292. Cladogram of the members of the mexicanus group 369

293. Cladogram of the members of the briggsi group 373

vm LIST OF MAPS

MAP Page

1. Distribution of schizomids of the cubanicus group 54

2. Distribution of schizomids of the simonis group . 54

3. Distribution of protoschizomids 168

4. Distribution of schizomids of the pecki and goodnightorum groups 168

5. Distribution of schizomids of the mexicanus group . 168

6. Distribution of schizomids of the briggsi group . . 132

7. Distribution of schizomids of the brasiliensis group 132

8. Evolutionary zoogeography of the cubanicus group . 381

9-11. Evolutionary zoogeography of the brasiliensis group 386

12-14. Evolutionary zoogeography of the mexicanus group . 392

15-17. Evolutionary zoogeography of the briggsi group . . 399

IX CHAPTER I

INTRODUCTION

The purposes of this research were: (1) to provide a classification of generic and familial taxa of the arachnid order

Schizomida on a world-wide basis, including a list of all species and a bibliography containing the entire primary literature; (2) to describe and classify the complete New World fauna as far as available material would allow; (3) to perform phylogenetic studies on the

World genera and familial taxa, and New World species; and (4) to analyze the zoogeographical relationships of taxa wherever evidence allowed. Phylogenetic and taxonomic considerations and results are

highly interdependent and were developed simultaneously, but are discussed separately herein.

The order Schizomida, like a few other minor arachnid orders, encompasses a manageable number of species of surprisingly few adaptive types. Though morphologically conservative, they have a

nearly complete pan-tropical distribution and are often abundant, and hence probably ecologically very important in certain areas.

These arachnids have received little attention, however, aside from

descriptive works, since the first was described in 1872. The first

critical study of the known species was that of Hansen and Sorensen

(1905) and aside from the taxonomy in this paper the anatomical

information is still quite useful. Lawrence (1969) produced a

revision of African fauna, which stands as the only regional

treatment. Millot (1949) gave the only detailed account, and Savory

1 (1964) gave a general account of all aspects of schizomid biology.

The works of Gravely (1915), Rowland (1972), Beck (1968a, 1968b), and Sturm (1958, 1973) offer all the information on behavior. The only major works on descriptive internal anatomy are those of Borner

(1902a, 1902b, 1904), Modder (1960), and Rowland (in preparation).

Post embryonic development of most schizomids probably consists of five instars over a period of two or three years. The nymphal instars resemble the adults but are smaller and lack, in varying degrees, the cuticular sclerotization of the adults. The sexual differences manifest in the flagellum, pedipalps and primary sexual organs occur at the last molt, although penultimate males are sometimes recognizable by the shape of the flagellum. The general breeding habits of schizomids were originally assumed to be similar to that of the uropygids. Subsequently, Hansen and Sorensen (1905) described a spermatophore attached to the venter of a female, confirming a similarity. Sturm (1958) published an account of his observations of the entire courtship and sperm transfer. Later

Sturm and Kraus (1973) and Beck (1967) added further behavioral data.

My personal observations on mating behavior concur with these sources.

After a few minutes of preliminary, mainly tactile, courtship, the male presents his flagellum to the female, which she envelops with the pedipalps and perhaps chelicerae. This action precedes the male's forward advancement which pulls the female over a spermatophore; the female then lowers herself onto it. In a few minutes the couple breaks but may remain in close proximity for some time. The spermatophore remains attached to the female for an unknown time but is expelled before oviposition. Oviposition has been documented only by Gravely (1915) and Rowland (1972a). They reported that females, kept in small terraria, constructed earthen chambers and that eggs were oviposited and attached to the venter of the parent. In the latter case young had emerged and were active in about three months.

Some species, contrary to the above observations, are known to carry eggs while roaming at large (R. W. Mitchell, J. R. Reddell, personal conmuni cation).

Schizomids are extremely sensitive to dessication and are most frequently found under rocks in mesic environments and where ample leaf litter is available to support an adequate arthropod fauna upon which they presumably prey. The sifting of deep, moist leaf litter may also produce schizomids, but generally not in the numbers sometimes encountered when turning over partially buried objects in moist, undisturbed soil. Two reports of sweeping schizomids from brush come from C. J. Goodnight and S. B. Peck (personal communications), which gives evidence that very much is still to be learned about the habits of this group of arachnids.

Anatomically, schizomidans may be considered the most generalized of all arachnids. The body consists of three major regions: prosoma, opisthosoma, and telson, or flagellum. The prosoma is dorsally divided into three transverse regions, each with one or two plates. The carapace, or propeltidium, overlies the first section, the paired mesopeltidia overlie the second section, and the entire or paired metapeltidium overlies the third section. Ventrally, the prosoma is composed of the five pairs of coxae and an anterior and posterior sternum. The opisthosoma is covered dorsally by nine terga and ventrally by nine sterna. The last three abdominal segments are not divided into terga and sterna, but are entire and compose the pygydial segments, that is, segments X-XII. The flagellum inserts on the posterior face of the last abdominal segment. The appendages of the prosoma are the chelicerae of two articles, the pedipalps of six articles, and legs I-IV each composed of seven articles. The tarsus of leg I is composed of six tarsomeres, the other legs of three tarsomeres. Leg I is not ambulatory, but functions as a tactile organ. Legs II-IV are ambulatory, the fourth always being the longest, the third always the shortest. The fourth

leg differs nost noticably in the great expansion and muscular development of the femur in most species. The central nervous system

is composed of a superesophageal ganglionic mass and a subesophageal ganglionic mass formed by the union of post-cheliceral pedal ganglia and the first four abdominal ganglia. An abdominal ganglionic mass

is located in abdominal segments III and IV and is composed of eight neuromeres. The breathing system is composed of one pair of book

lungs, communicating outside by a spiracle on the posterolateral margin of each side of abdominal segment II. The heart is limited

to the second to seventh abdominal segments, and has five pairs of ostia. The cephalic end of the heart leads into a dorsal blood

vessel, which extends to the subesophageal ganglionic mass; however. no abdominal arterial system is present. The digestive system is provided with two prosomal midgut diverticulae and a few large opisthosomal diverticulae. The anus opens just below the insertion of the flagellum. The coxal glands open at the base of leg I, but originate in the sixth prosomal segment. No definite saccule exists.

Defensive organs, as in the uropygids, consist of a pair of anal glands which produce and emit an acetic acid spray. The male's genital system is composed of four major parts: the paired testes, the dorsal tube system, the vasa seminala, and the gonopore. The female's genital system is simpler. It is composed of two ovaries which are fused into a single mass provided with two short oviducts.

These lead into a simple uterus, which in turn opens directly to the outside.

Only 170 species, including those described herein, of a probably vastly larger World fauna are known. This study of Mexican species shows that there are few wide ranging species, most being highly localized. They lack any secondary means of travel, such as phoresy, ballooning, resistant eggs, parasitism, or other means employed by arachnids, and are thus relatively non-vagile. It appears, for example, that only one New World species has a distribution that is continuous for more than one hundred miles.

Intrageneric sympatry in epigean habitats occurs, as far as is known, only in tropical areas. Sympatry in subtropical areas is limited to caves, and is unknown in temperate areas. Members of this order have sometimes been combined with the larger whip scorpions, the Uropygida, into a single order. Most

American and English tastes tend toward separation of the two, though nothing of great consequence is achieved in either view. Most importantly, no doubt has been raised that the members of the order

Schizomida represent a monophyletic assemblage with unique evolutionary tendencies, and are a result of an ancient dichotomy from common Uropygida-Schizomida stock. The age of dichotomy lies hidden in antiquity, but must considerably predate the appearance of fossil uropygids in the Carboniferous, which are practically indistinguishable from modern forms. Moreover, the schizomidans form a unique assemblage of species which most probably will never form a morphological continuum with the uropygids through discovery of extant organisms. Rowland (in preparation) reviews the comparative anatomy of these groups.

These animals are commonly called "schizomids," but other

English common names are "tartarides" and "micro-whip scorpions."

In this work I have used, for purposes of clarity, the word "schizomids" only in a restricted sense, namely to designate members of the family

Schizomidae as opposed to the members of the Protoschizomidae. The collective term for members of both families, hence the order, is here "schizomidans." It is not my intent nor my wish to change the vernacular, but to minimize confusion in this text. The Bibliography contains all of the primary literature on schizomidans and additional works not concerning schizomidans,

The latter titles bear an asterisk. Fig. 1.--Above, male and below, female, Agastoschizomus lucifer, from Sotano de Yerbaniz, San Luis Potosi.

Fig. 2.—Above, male (left) and fanale (right) Schizomus pentapeltis from near Palm Springs, California, during courtship.

Below, male S. mexicanus from Sotano de la Tinaja, San Luis Potosi.

Figs. 3-6.--Generalized schizomidan anatomy: 3, dorsal view of male; 4, lateral view, right pedipalp; 5, lateral view, right leg I;

6, lateral view, right leg II. A, apical carapacal setae; E, eyespot;

D, dorsal carapacal setae; CA, carapace; MS, mesopeltidial plate;

MT, metapeltidial plate (in the split condition); Tl, abdominal

tergum I; T2, abdominal tergum II; T9, abdominal tergum IX; AP,

posterodorsal abdominal process; FL, flagellum; T, trochanter; F,

femur; P, patella; Tl, tibia; TB, tarsus-basitarsus; S, tarsal-

basitarsal spur; C, claw; B, basitarsus; TA, tarsus. Scale: 1 cm =

.25 mm. 13 Figs. 7-13.—Generalized schizomidan anatomy: 7, 8, mesal views of right chelicerae: 7, Schizomus sp.; 8, Agastoschizomus

lucifer; 9, mesal view of moveable finger of right chelicerae of

Megaschizomus mossambicus; 10-13, lateral view of left leg IV:

10, Schizomus sp.; 11, Megaschizomus mossambicus; 12, Protoschizomus

pachypalpus; 13, Agastoschizomus lucifer. 8, brush; F, fixed finger;

S, serrula. Scale: Fig. 7, 1 cm = .12 mm; Figs. 8, 9, 1 cm = .25 mm;

Figs. 10, 12, 1 cm = 3 mm; Figs. 11, 13, 1 cm = .8 mm. 15 -y?^ CHAPTER II

MATERIALS AND METHODS

This study was based on examination of approximately 1,000 specimens of schizomidans from the New and Old World. The vast majority of these were from the New World and formed the basis of my revisionary work for that area. A small part of these collections was private, but will be deposited in various permanent collections.

Public collections were borrowed from the following institutions.

Curators are also acknowledged and abbreviations for institutions are given where I have indicated deposition of material. American Museum of Natural History, New York (AMNH), J. A. L. Cooke, N. I. Platnick;

Bernice P. Bishop Museum, Honolulu, Hawaii, J. Tenorio; British

Museum (Natural History), London (BMNH), K. H. Hyatt, F- R. Wanless;

California Academy of Sciences, San Francisco, California (CAS), R.

X. Schick; Institut royal des Sciences naturelles de Belgique,

Brussells (ISB), N. Leleup; Institute of Zoology, Uppsala (IZU), A.

Holme; Instituto de Zoologia, Academia de Ciencias, La Habana, Cuba

(IZC), L. F- de Armas; Los Angeles County Museum, Los Angeles,

California (LACM), C. L. Hogue; Musee National d'Histoire Naturelle,

Paris (MNHN), M. Vachon; Museum of Comparative Zoology, Harvard

University, Cambridge, Massachusetts (MCZ), H. W. Levi; Natal Museum,

Pietermaritzburg, South Africa, B. H. Lamoral; Naturhistoriska

Riksmuseet, Stockholm, Sweden (NRS), T. Kronestedt; Naturhistorisches

Museum, Wien, Austria, M. Beier, J. Gruber; Natur-Museum und

Forschungs-Institut Senckenburg, Frankfurt, West Germany (SMF), M.

16 17

Grasshoff; San Diego Natural History Museum, San Diego, California,

A. Ross; South Australia Museum, R. V. Southcott; Koninklijk Museum

Voor Midden Africa (KMMA), P. L. G. Benoit, N. Leleup; The Museum,

Texas Tech University, Lubbock, Texas (TTU), R. W. Mitchell, J. R.

Reddell; Universitetets Zoologiske Museum, Copenhagen, Denmark

(UZMK), S. L. Tuxen; United States National Museum, Washington, D. C.

(USNM), R. Crabill. Individuals contributing private collections are as follows: T. S. Briggs, C. J. Goodnight, K. Hom, W. Icenogle,

B. J. Kaston, W. Lum, W. B. Muchmore, S. B. Peck, W. B. Peck, B.

Phillips, W. B. Shear, I. Shimojana, M. E. Thompson.

Observational methods. An M-5 Wild dissecting microscope was used for general observations, measurements, and drawings of all anatomical parts other than the female's spermathecae. Measurements were made at lOOX with lOX oculars and a 2X objective boost.

Drawings of the pedipalps and whole views were accomplished by use of the M-5 Zeichentubus, a camera lucida. Other drawings were made by the grid method using a Wild, 20 square grid micrometer and one half inch square grid paper. For work with the dissecting scope specimens were observed while in 80% ethyl alcohol in a Syracuse watch glass using fine, clean silica sand as a substrate. All specimens were stored in 80% ethyl alcohol, unless previously preserved otherwise by the lending institution.

To observe and illustrate the female's spermathecae it was necessary to dissect the second (genital) and third abdominal sterna by use of instruments fashioned from minute pins. The female's 18 spermathecal structures are closely associated with the internal surface of sternum II and come free with it. The entire piece, consisting of the spermathecae, sternum II and III, the book-lungs, and various muscular tissue, was then carefully cleaned to remove extraneous materials. The cleaned piece was then immersed in three drops of clove oil on a slide and allowed to heat for three minutes at 75 °C. A cover slip was then applied to flatten the entire

structure. The details of the spermathecae were then drawn using an

AO Spencer phase-contrast binocular microscope using an ocular micrometer at 200X and one-fifth inch square grid paper. The

spermathecae and associated pieces were then washed in several changes

of alcohol and stored in small vials with the remainder of the

specimen.

Descriptive statistics. Accompanying each description is a

set of descriptive statistics given in tabular form which I provide

for all species. In the case of S^. portoricensis the range, mean,

standard deviation, and coefficient of variation are given for

several populations. All other species are represented by the range

of measurements only. The sample size of most species is so low

(usually two or three specimens) that a comprehensive set of

statistics measuring variation is impossible. The statistics are

intended only for general comparisons and species recognition.

Relative lengths given in ratios of certain pairs of anatomical parts

are used only as a convenient method of indicating rough body

proportions. The variates are as follows: 1, length of carapace; 19

2, length of flagellum; 3, width of flagellum; 4, length of femur I;

5, length of patella I; 6, length of tibia I; 7, length of tarsus-

basitarsus I; 8, length of femur II; 9, length of patella II; 10,

length of tibia II; 11, length of basitarsus II; 12, length of

femur III; 13, length of patella III; 14, length of tibia III; 15,

length of basitarsus III; 16, length of femur IV; 17, length of

patella IV; 18, length of tibia IV; 19, length of basitarsus IV.

Criteria for taxonomic judgments. Since very few specimens of most species were available for study, and since most species are

known from a single locality or even a single collection, almost no

understanding of the morphological and geographical limits of species

and populations is forthcoming. Thus, the biological species concept,

as in most taxonomic works, has no relevance. I have used the

classical, arbitrary morphological species concept because no other

definition is workable within this poorly known group.

In cases where morphologically distinct, yet closely related,

forms occur without known morphological or geographical continuity,

they were given separate specific status. This was done because it

is probably best to risk indicating spurious variation within a taxon

than to risk erroneously hiding it. I have not used the subspecies

category since the materials available to me do not facilitate such

judgments. Species groups were given loose and arbitrary limits based

mainly on morphological, but also on distributional, data. The use

of species groups and complexes serves well in foundation of

polythetic categories of monophyletic or at least paraphyletic 20 origins, and serve equally well as subgenera, but have the advantage

of not carrying with them any of the legalistic problems of formal

taxonomy.

I defend my use of generic, subfamilial, and familial

categories only in that in my judgment they reflect discrete phyletic

entities which are typified by correlated sets of characters. These

taxa will probably not show continuity with contemporary forms of

equal rank, due to long and unique evolutionary histories in which

intermediate forms have become extinct or have themselves changed

by evolutionary gradualism. My philosophies concerning the

construction of hierarchies and development of the taxa, are

discussed in the section on phylogeny.

The characters upon which taxonomic and phylogenetic judgments

are made are discussed, when necessary, in the section on phylogeny.

Recognition of taxa. Recognition of the various taxa treated

in this work is facilitated by use of comparisons tables. Keys seem

inappropriate for use in this study for various reasons. It is my

estimation that future efforts will produce considerable numbers of

new species within the groups delineated here. Keys are best

employed where the fauna is completely, or nearly completely, known

and where the chance of encountering undescribed taxa is relatively

small. Thus, I have provided only comparisons tables which are

equally as useful as keys in distinguishing described taxa, but which

are perhaps not as expedient. Comparisons tables have the added 21 advantage of presenting considerably more contrasting information than a key, and facilitate placing undescribed taxa.

I do not expect that any large species groups, genera, or familial taxa will be discovered in the New World; therefore, keys may be fairly safely used for taxa above the species level. Again

I have refrained from using keys, but for the reason that there are so few taxa to recognize. The two family group and three generic taxa are described only in contrasting characters and are essentially key couplets in themselves.

Upon studying the criteria for placement of taxa into family categories the user will, in the case of the family Schizomidae, refer to the comparisons table for species groups. This will direct him to the appropriate species group under which he will inspect the comparisons table for those species. Augmenting the table with examination of the illustrations of appropriate taxa, the user will be able not only to identify known species, but may also arrive at some idea of the relationships of undescribed taxa to species treated herein. CHAPTER III

CLASSIFICATION

Systematic History. Currently there are seven familial and

13 generic names applied to extant organisms and two familial and three generic names applied to extinct organisms within the order

Schizomida. The main purpose of this section of this chapter is to examine the applicability and usage of the names of extant taxa.

Calcitronidae and its two monotypic genera are wholly extinct and known only from Pliocene calcite deposits in Arizona. The works of Petrunkevitch (1945a, 1955), Pierce (1950, 1951), and Rowland

(1973b) provide all the information available on these taxa.

Calcoschizomus latisternum Pierce, is apparently also extinct, known only from the same deposits as the Pliocene calcitronids, but has been included in the Schizomidae. I have examined all of these types, but I have not been able to confirm many of the observations made by the original authors. These specimens are interesting, but of limited use since critical observations cannot be made. I have chosen not to integrate the taxonomy of the extinct and extant taxa here. In

Rowland (1973b) I included the fossil material and gave diagnoses which were repeated from the original sources. I have little hope now that these fossils were accurately described, or that they can ever be properly interpreted.

Cambridge (1872) described the first schizomid and created for it the family Tartarides, genus Nyctalops, and trivial name crassicaudata. In this work he also described j^. tennuicaudata. The

22 23 latter name was subsequently moved to synonymy of Pocock (1900) on the basis that N^. tennuicaudata was actually the female of H_. crassicaudata. Evidence is convincing that this revision is sound, although Pocock

(1893) had earlier assigned N^. tennuicaudata to another genus. Cook

(1899), by authority of first revisor, designated crassicaudata as type of the genus.

Previous to Pocock's work Thorell (1888) elevated Cambridge's family Tartarides to tribal status (tribal rank was then superior to family rank) and introduced the new family name Schizonotoidae. He proved that the name Nyctalops is a junior homonym and had been previously occupied (Hagler, 1832). In its place he provided

Schizonotus, the nominate genus of Schizonotoidae. The spelling of the latter name was emended to Schizonotidae by Pocock (1893). The work of Cook (1899) shows that the name Schizonotus is also a junior homonym, being preoccupied (Ratzeburg, 1852). In its place he used the name Schizomus. For the family name Schizonotidae, which was based on the junior homonym, he substituted the new name Hubbardiidae, which contained a new nominate genus Hubbardia; however the type genus remained Schizomus.

Thorell (1899) erected the new genus Tripeltis on the basis of two new species, J_. grassii and X- cambridgei, the former being originally designated as type species of the genus. The work of

Cook (1899), however, shows that Tripeltis is a junior homonym, being preoccupied (Cope, 1886). For Tripeltis he provided the name Triplomus.

Cook's revision does not stand, however, since Kraepelin (1899) 24 offered the name Trithyreus for Tripeltis for similar reasons, just one month before Cook.

Cook (1899) described the new genus and species Hubbardia pentapeltis; however, Hansen and Sorensen in 1905 synonymized this genus with Trithyreus. Cook (1899) also mentioned a new animal which he had presented at a meeting of the Entomological Society of Washington two years previously as Artacarus liberiensis. A printed description of this species never appeared under his authorship and, consequently, it was treated as a nomen nudum by Kraepelin (1897) and Hansen and

Sorensen (1905). Kraus (1960) located the original type series and described the species and placed it in Schizomus.

The original classification of Cambridge (1872) included the family name Tartarides; there was no nominal genus, but the type genus

Nyctalops was fixed by monotypy. Thorell (1888) elevated Tartarides to tribal status, but did not use a nominate family name. He used the new name Schizonotoidae, which is invalid because the nominate genus Schizonotus was later shown to be a junior homonym. Family

Hubbardiidae, which Cook (1899) offered to replace Schizonotidae was not allowed because its nominate type genus was later decided to be a junior subjective synonym of Schizomus. The family name Schizomidae has been erroneously attributed to Chamberlin (1922). Hansen and

Sorensen (1905) offered the name Schizomoidae to replace Hubbardiidae.

The original spelling was emended to Schizomidae by Gravely (1915).

The family name Tartarides should have been conserved from the start by Thorell and given its proper suffix, but since the name has been 25 out of general usage, the name Schizomidae should be used (Rowland,

1973b). In the latter work I introduced the subfamily Megaschizominae for reception of Megaschizomus Lawrence, 1969 and Agastoschizomus

Rowland, 1971. In Rowland (1975) I introduced Protoschizomidae for

reception of Agastoschizomus and Protoschizomus Rowland, 1975, and revised Megaschizominae, leaving it monobasic within the Schizomidae.

In the same publication that Chamberlin made his revision of family names he also described a new genus and species from Puerto

Rico. This genus, Stenochrus, was appropriately synonymized with

Schizomus in Rowland (1973b) and S^. portoricensis, in fact, later proved to be synonymous with some other rather well known species.

Lawrence (1969) synonymized African schizomids of the genus

Schizomus and Trithyreus. Unfortunately, Lawrence apparently missed the earlier revision of a similar nature by Mello-Leitao (1931).

Mello-Leitao reasoned that the two genera should be reduced to subgenera and that only the genus Schizomus should stand. While I agree with Lawrence in uniting the two genera, it seems he, like

Mello-Leitao, picked the younger generic name to survive. Trithyreus

Kraepelin dates to March, 1899, whereas Schizomus Cook dates to April,

1899.

After considering the Trithyreus-Schizomus dilemma for a few years I have decided to let Trithyreus stand. I do not recognize the genus as it was originally, or even as it recently has been conceived.

The split metapeltidium may occur erratically within a single species, or species group, and should never be used by itself to distinguish 25 out of general usage, the name Schizomidae should be used (Rowland,

1973b). In the latter work I introduced the subfamily Megaschizominae for reception of Megaschizomus Lawrence, 1969 and Agastoschizomus

Rowland, 1971. In Rowland (1975) I introduced Protoschizomidae for reception of Agastoschizomus and Protoschizomus Rowland, 1975, and revised Megaschizominae, leaving it monobasic within the Schizomidae.

In the same publication that Chamberlin made his revision of family names he also described a new genus and species from Puerto

Rico. This genus, Stenochrus, was appropriately synonymized with

Schizomus in Rowland (1973b) and S^. portoricensis, in fact, later proved to be synonymous with some other rather well known species.

Lawrence (1969) synonymized African schizomids of the genus

Schizomus and Trithyreus. Unfortunately, Lawrence apparently missed the earlier revision of a similar nature by Mello-Leitao (1931).

Mello-Leitao reasoned that the two genera should be reduced to subgenera and that only the genus Schizomus should stand. While I agree with Lawrence in uniting the two genera, it seems he, like

Mello-Leitao, picked the younger generic name to survive. Trithyreus

Kraepelin dates to March, 1899, whereas Schizomus Cook dates to April,

1899.

After considering the Trithyreus-Schizomus dilemma for a few years I have decided to let Trithyreus stand. I do not recognize the genus as it was originally, or even as it recently has been conceived.

The split metapeltidium may occur erratically within a single species, or species group, and should never be used by itself to distinguish 26 any taxon. Since the type and only known specimen of X- grassii, the species which originally was the basis for creation of Trithyreus, has apparently been lost in a flood, no subjective evaluation can be made until the species is recollected. It is possible, however, that this species may best be contained in a genus apart from Schizomus, because

Hansen and Sorensen (1905) made mention of several characters which they deemed ^ery distinctive.

Whatever the situation with X- grassii, I am at this point reluctant to synonymize Trithyreus and Schizomus, but I limit my use of the former genus for its type species, and remove all other species once assigned within Trithyreus to Schizomus. Until a study of X- grassii is made no further species should be assigned to Trithyreus.

For the sake of clarity I have provided a list of all schizomids, within their proper genera, as now conceived.

Order Schizomida

Protoschizomidae Rowland, 1975

Protoschizomidae Rowland, 1975:1.

Description.--Carapace without eyespots; mesopeltidia large, separated by about 1/3 to 1/10 their greatest dimension; metapeltidium entire or divided. Abdomen with eight pairs of dorsoventral muscles.

Flagellum segmented in females. Cheliceral serrula absent, but represented by a row of blunt, nearly hemispherical knobs; no row of closely situated setae at base of fixed digit (brush). Two teeth on 27 fixed digit, basal one arising at an angle from ventral surface of basal segment. Ratio of pedipalpal claw length to dorsal length of tarsus-basitarsus 1:.5 or 1:1; basitarsal spurs symmetrically placed, long, about 1/3 to 1/2 dorsal length of tarsus-basitarsus; "true spines" present. Trochanter IV about 2.2 times longer than wide; femur IV from about three to five times longer than deep.

Di stri buti on.--Mexi co.

Protoschizomus Rowland, 1975

Protoschizomus Rowland, 1975:2. Type: Agastoschizomus pachypalpus

Rowland, 1973 (orig. desig.).

Description.--Carapace about one millimeter in length.

Mesopeltidial plates separated by 1/3 their length; greatest length to width ratio of metapeltidial plates about 1.3:1. Abdominal sterna

IV to VII about six times wider than long. Pedipalpal claw shorter than dorsal length of tarsus-basitarsus. Femur IV about three times longer than deep.

Distribution.--Tamaulipas and Colima, Mexico.

P.. pachypalpus (Rowland), 1973 - Tamaulipas

P.. occidental's Rowland, 1975 - Colima

Agastoschizomus Rowland, 1971

Agastoschizomus Rowland. 1971:13. Type: Agastoschizomus lucifer

Rowland, 1971 (monotypy); Rowland, 1973a:202; Rowland, 1975:5. 28

Agastoschizomus Rowland, 1971

Agastoschizomus Rowland, 1971:13. Type: Agastoschizomus lucifer

Rowland, 1971 (monotypy); Rowland, 1973a:202; Rowland, 1975:5,

Description.--Carapacal length about two to three millimeters in length. Mesopeltidial plates separated by 1/10 their greatest dimension; greatest length to greatest width ratio of metapeltidial plates about 1:1. Abdominal sterna IV to VII about three times wider than long. Pedipalpal claw longer than dorsal length of tarsus-basitarsus, ratio of about 1.1:1; tarsal-basitarsal spurs about 1/2 dorsal length of tarsus-basitarsus. Femur IV about five times longer than deep.

Distribution.--San Luis Potosi, Mexico.

A. lucifer Rowland, 1971 - Tamaulipas, San Luis Potosi

A. huitzmolotitlensis Rowland, 1975 - San Luis Potosi

Family Schizomidae Hansen and Sorensen, 1905

Tartarides Cambridge, 1872:410 (nom. obi.).

Schizonotidae Thorell, 1888:358 [nom. correct. Pocock, 1893 (ex

Schizonotoidae Thorell, 1888, nom. imperf.)] (name based

on junior homonym).

Hubbardiidae Cook, 1899:429 (nom. obi.) (name based on junior

subjective synonym).

Schizomidae Hansen and Sorensen, 1905:4 [nom. correct. Gravely, 1915

(ex Schizomoidae Hansen and Sorensen, 1905, nom. imperf.)] 29

Schizomidae Chamberlin, 1922:11 (jun. prim. hom,).

Type: Nyctalops crassicaudatus Cambridge, 1872.

Description.--Carapace with or without eyespots, mesopeltidia small, separated by about 1/2 to 2/3 their greatest dimension; metapeltidium entire or divided. Abdomen with seven pairs of dorsoventral muscles. Flagellum segmented or unsegmented in females.

Cheliceral serrula present, a row of closely situated setae also present at base of fixed digit (brush). Three or five to nine teeth present on fixed digit, basal one arising gradually from ventral surface of basal segment. Ratio of pedipalpal claw length to tarsal-basitarsal length from about 1:4 to 1:1.2; tarsal-basitarsal spurs asymmetrically placed, medium to short, about 1/4 to 1/5 dorsal length of tarsus-basitarsus; "true spines" present or absent.

Trochanter IV about 1.4 or 2.2 times longer than wide, femur IV from about 2.6 to 4.1 times longer than deep.

Distribution.--Present in all tropical and certain subtropical and temperate areas.

Schizominae Hansen and Sorensen, 1905

Schizomidae, Hansen and Sorensen, 1905:4.

Schizominae, Rowland, 1973a:200 [nom. transl. Rowland, 1973a (ex

Schizomidae Hansen and Sorensen, 1905)]; Rowland, 1975:11.

Type: Nyctalops crassicaudata Cambridge, 1872 30

Description.--Carapace with or without eyespots; mesopeltidia divided by about equal to their greatest dimension; metapeltidium entire or divided. Flagellum unsegmented in females. Fixed digit of chelicerae with five to nine teeth. Ratio of pedipalpal claw to tarsal-basitarsal length about 1:4 to 1:1.2; tarsal-basitarsal spurs asymmetrically placed, short, about 1/5 dorsal length of tarsus-basitarsus; "true spines" absent. Trochanter IV about 1.4 times longer than wide, femur IV about 2.6 times longer than deep.

Distribution.--Same as for the family.

Schizomus Cook, 1899

Nyctalops Cambridge, 1872:410. (jun. hom.) Type: N^. crassicaudata

(sub. des.. Cook, 1899).

Schizonotus Thorell, 1888:358. (jun. hom.) [nom. subst. pro Nyctalops

(non Nyctalops Wagler, 1832)]; Kraepelin, 1897:50; Kraepelin,

1899:233.

Tripeltis Thorell, 1889:554 (jun. hom.). Type: X- q'^assii (orig. des.)

Trithyreus Kraepelin, 1899:234 [nom. subst. pro Tripeltis (non

Tripeltis Cope, 1886)] in part.

Triplomus Cook, 1899:250. (jun. obj. syn.) [nom. subst. pro Tripeltis

(non Tripeltis Cope, 1886)

Hubbardia Cook, 1899:250. (jun. subj. syn.). Type: H.. pentapeltis

Cook (monotypy).

Artacarus Cook. 1899:254. (jun. subj. syn.) Type: A. liberiensis

Cook (monotypy). 31

Schizomus Cook, 1899:249 [nom. subst, pro Schizonotus (non Schizonotus

Ratzeburg, 1852).

Stenochrus Chamberlin, 1922:11 (jun. subj. syn.) Type: S^,

portoricensis Chamberlin (monotypy),

Heteroschizomus Rowland, 1973:1 (jun, subj, syn,) Type: h[,

goodnightorum Rowland (monotypy), NEW SYNONYMY

Description,--Same as that for the subfamily.

Distribution,--Same as that for the family.

S^. crassicaudatus (Cambridge), 1872 - Ceylon

S^. cambridgii (Thorell), 1889 - Burma NEW COMBINATION

S^. liberiensis (Cook), 1897 - Liberia

S^. pentapeltis (Cook), 1899 - California

S^. suboculatus (Pocock), 1900 - Ceylon

S^. dispar Hansen and Sorensen, 1905 - Martinique

S^. flavescens Hansen and Sorensen, 1905 - Venezuela

S^. insignis Hansen and Sorensen, 1905 - Martinique

^. simonis Hansen and Sorensen, 1905 - Venezuela i- ^atipes Hansen and Sorensen, 1905 - Seychelles i- africanus (Hansen and Sorensen), 1905 - Sierra Leone i- claviger (Hansen and Sorensen), 1905 - Malaysia i. luzonicus (Hansen and Sorensen), 1905 - Philippines i. modestus (Hansen and Sorensen), 1905 - New Guinea i, procerus (Hansen and Sorensen), 1905 - Malaysia

S_. siamensis (Hansen and Sorensen), 1905 - Thailand 32

S^. montanus Hansen, 1910 - Tanzania

S^. sauteri Kraepelin, 1911 - Vietnam

S^. bagnallii (Jackson), 1911 - England (introduced)

S^. lunatus (Gravely), 1911 - India

S^. peradeniyensis (Gravely), 1911 - Ceylon

S^. vittatus (Gravely), 1911 - Ceylon

S^. cavernicola Gravely, 1912 - Burma

S.. greeni (Gravely), 1912 - Ceylon NEW COMBINATION

S^. kharagpurensis (Gravely), 1912 - India

S^. similis Hirst, 1913 - Seychelles

S^. buxtoni Gravely, 1915 - Ceylon

S^. perplexus Gravely, 1915 - Ceylon

S^. breviCauda (Hansen), 1921 - Portuguese Guinea

S^. parvus (Hansen), 1921 - Equitorial Guinea

S^. portoricensis (Chamberlin), 1922 - New World tropics

S^. guatemalensis Chamberlin, 1922 - Guatemala

S^. sijuensis Gravely, 1924 - India

S^. hanseni (Mello-Leitao), 1931 - Zanzibar

S^. antilus Hilton, 1933 - Cuba

S^. ghesquierei (Giltay), 1935 - Zaire i. wessoni (Chamberlin), 1939 - Arizona NEW COMBINATION i- davisi Gertsch, 1940 - Tamaulipas i- mulaiki Gertsch, 1940 - Texas i. centralis Gertsch, 1941 - Panama i- sawadai Kishida, 1943 - Bonin Islands (Japan) 33

S. peteloti (Remy), 1945 - Vietnam NEW COMBINATION S^. schoutedeni (Roewer), 1954 - Zaire S^. formicoides Fernando, 1957 - Ceylon S_. belkini (McDonald and Hogue), 1957 - California NEW COMBINATION S^. cumbalensis (Kraus), 1957 - Columbia NEW COMBINATION S.. sturmi (Kraus), 1957 - Columbia NEW COMBINATION S.. macarensis (Kraus), 1957 - Columbia NEW COMBINATION S^. machadoi Lawrence, 1958 - Angola S^. gladiator Remy, 1961 - Surinam S^. surinamensis Remy, 1961 - Surinam S^. vanderdrifti Remy, 1961 - Surinam ^. borregoensis (Briggs and Hom), 1966 - California NEW COMBINATION S. brasiliensis (Kraus), 1967 - Brazil NEW COMBINATION S^. vadoni Lawrence, 1969 - Madagascar S^. mediocriter Lawrence, 1969 - Tanzania S^. madagassus Lawrence, 1969 - Madagascar S^. tenuipes Lawrence, 1969 - Rodrigues Island (Mauritius) i. virescens Lawrence, 1969 - Rodrigues Island (Mauritius) S^, pauliani Lawrence, 1969 - Madagascar S^. vinsoni Lawrence, 1969 - Mauritius i- lemzi Lawrence, 1969 - Reunion Island i. milloti Lawrence, 1969 - Madagascar S^, benoiti Lawrence, 1969 - Somalia S^. nidicolous Lawrence, 1969 - Zaire i- mexicanus Rowland, 1971 - Tamaulipas 34

!• mitchelli Rowland, 1971 - Tamaulipas

S^. cookei Rowland, 1971 - San Luis Potosi

S^, reddelli Rowland, 1971 - Tamaulipas

S^. joshuensis (Rowland), 1971 - California NEW COMBINATION

S., briggsi (Rowland), 1972 - California NEW COMBINATION

S^. shoshonensis (Briggs and Hom), 1972 - California NEW COMBINATION

S.. goodnightorum (Rowland), 1973 - Yucatan NEW COMBINATION

S^, orthoplax Rowland, 1973 - Chiapas

S^, bartolo Rowland, 1973 - Nuevo Leon

S^. firstmani Rowland, 1973 - Veracruz

S.- pecki Rowland, 1973 - Tabasco

S^. sbordonii Brignoli, 1973 - Veracruz

S^. arganoi Brignoli, 1973 - Chiapas

S^. stewarti Rowland, 1973 - Oaxaca

S^, moisii Rowland, 1973 - Oaxaca

S^, lukensi Rowland, 1973 - Tamaulipas

S^. negreai Dumitresco, 1973 - Cuba

S^. rowlandi Dumitresco, 1973 - Cuba

S^. lanceolatus Rowland, 1975 - Veracruz i, trilobatus Rowland, 1975 - Tabasco

S^. pallidus Rowland, 1975 - Veracruz i. lacandonus Rowland, 1975 - Chiapas i- infernal is Rowland, 1975 - Chiapas i- cubanicus, new species - Cuba

S^. cubanicoides, new species - Costa Rica 35

S^. brevi patellatus, new species - Haiti

S_. longipalpus, new species - Haiti

S^. primibiconourus, new species - Jamaica

S^. cousinensis, new species - Jamaica

S^. subiconourus, new species - Cuba

S^. desecheo, new species - Puerto Rico

S^. monensis, new species - Puerto Rico

S^. biconourus, new species - Cuba

S^. peckorum, new species - Jamaica

S^. viridis, new species - Jamaica

S^. drakos, new species - British Guiana

!• trinidanus, new species - Trinidad

S^. acrocaudatus, new species - Trinidad

§.' tobago, new species - Tobago

S. mumai, new species - Costa Rica

S^. cuenca, new species - Ecuador i- pallipatellatus, new species - Costa Rica

S^. silvino, new species - Guatemala i- armasi, new species - Cuba

S^. troglobius, new species - Jamaica

Trithyreus Kraepelin, 1899

Tripeltis Thorell. 1889:554, in part.

Trithyreus Kraepelin, 1899:234, in part; Hansen and Sorensen, 1905:34,

64, in part. 36

Description.--Same as that for Trithyreus grassii (Thorell),

1889.

Di stri buti on.--Tei nzo, Burma.

X- gi^assii (Thorell), 1889 - Burma

Megaschizominae Rowland, 1973

Megaschizominae Rowland, 1973b:202; Rowland, 1975:20.

Description.--Carapace without eyespots; mesopeltidia separated by about 2/3 their greatest dimension; metapeltidium entire.

Flagellum segmented in females, Chelicerae with three teeth on fixed digit. Ratio of pedipalpal claw length to dorsal tarsal-basitarsal length about 1:2; tarsal-basitarsal spurs slightly asymmetrical, medium sized, about 1/4 dorsal length of tarsus-basitarsus; "true spines" present. Trochanter IV about 2.2 times longer than wide, femur IV about 4,1 times longer than deep.

Distribution.--Southeastern Africa,

Megaschizomus Lawrence, 1969

Megaschizomus Lawrence, 1969:257, Type: Schizomus mossambicus

Lawrence, 1958 (orig. des.).

Description.--Same as that for the subfamily.

Distribution.--Same as that for the subfamily.

M. mossambicus (Lawrence), 1958 - Mozambique

M. zuluanus (Lawrence), 1947 - South Africa 37

The American Schizomida

The order Schizomida is represented in the New World by the two genera of the Protoschizomidae, which include only four species, and by the nominate subfamily and genus of the Schizomidae, which include the remainder of the 79 OTU's treated in this study. The only superspecific taxa not occurring in the New World are the

African subfamily Megaschizominae and the genus Trithyreus, which, as previously discussed, is only maintained on a nominal basis.

The genus Schizomus is by far the dominant group throughout the range of the order.

I have been able to make only a cursory examination of relatively little Old World material. The African species have been treated by Lawrence (1969), but the remainder of the Old World fauna remains largely untreated beyond original descriptive work.

The New World members of the genus Schizomus have been found to constitute several distinct lineages, six of which are here hypothesized to stem from an ancient New World stock and one of which seems to represent a more recent invading lineage from the Old

World. With the exception of four species, which I have not assigned to categories below that of genus, the New World Schizominae are placed in the following groups: the mainly Antillean cubanicus group, the middle and South American simonis group, the mainly middle and

South American brasiliensis group, the mainly Mexican mexicanus group, the Mexican and middle American pecki and goodnightorum groups, and the Californian and Arizonan briggsi group. Species within the above Table 1 .—Comparisons of the species groups of the American members of the genus Schizomus. For explanation of the characters see Phylogeny. 39

cuba­ brasil- mexl- CHARACTER simonis "::::,•; •:::;.; pecki 9°°^;;:^ briggsi nicus 1 ens Is canus

DORSAL 2-3 2-3 3-4 2-3 2-3 3-4 3-4 SETAE

METAPEL­ TIDIUM entire entire ^^lILo'entire " ent^i'e entire entire ^P!ientir< °*e "

brown or brown or brown or brown or (,rown brown '""Ow t"* COLOR green green green green orown green

SPERMA­ THECAE ML H>L H>L multiple

ART. FEM. FLAGELLUM

CARAPACE LENGTH .96-1.37 1.07-1.34 .91-1.48 .98-1.37 1.31-1.74 .89-1.42 1.18-1.52

ABDOMINAL none present none none none present ELONGATION present

ABDOMINAL present present present absent absent absent present PROCESS

PEDIPALPAL slight to slight to none to none to DIMORPHISM strong strong strong none none strong

long or SHAPE MALE bulbous long bulbous bulbous bulbous long FLAGELLUM bulbous 40 taxa have been been further grouped, where expedient, into complexes for purposes of discussion in the zoogeography section.

Table 1 presents the polythetic basis for each group and functions in taxa recognition. For the latter the user is directed to Contents, where location in the text of the treatment of each group is given.

For each of the following OTU's I have provided a reference to appropriate figures; tables; maps; a complete synonymic list, with corresponding bibliographic citations; the origin and disposition of all known type material; a description and further relevant statements. The measurements and illustrations are of the type material, unless otherwise stated.

Protoschizomus pachypalpus (Rowland), 1973

(Figs. 15, 18, 19; Table 2; Map 3)

Agastoschizomus pachypalpus Rowland, 1973:8-10,

Protoschizomus pachypalpus, Rowland, 1975:2-4,

Holotype.--An adult female, taken at Nacimiento del Rio Frio,

3 mi S Gomez Farias, Tamaulipas, Mexico, on 12 March 1969 by James

Reddell (AMNH, examined),

Paratypes,--Two adult females and one juvenile taken at

Arroyo Nacimiento del Rio Frio, Tamaulipas, Mexico, on 16 February

1970 by R. W, Mitchell (AMNH, examined); an adult female taken at

Gomez Farias, Tamaulipas, Mexico, on 6 January 1964 by J, Reddell, 41

D. McKenzie, and L, Manire (AMNH, examined); and one juvenile taken with the holotype (AMNH, examined),

Male,--An adult, taken 51,5 mi E Ciudad Victoria, Tamaulipas,

Mexico, on Highway 70, on 17 October 1972, by B. L. Firstman and

V, D. Roth (AMNH, examined).

Description.--Male. Color brownish. Carapace with five pairs of dorsal and two apical setae. Eyespots absent. Metapeltidium divided medially into two plates. Anterior sternum with 11 entire setae. Abdominal terga I to II with two setae. III to VII with four setae, VIII with 10 setae, IX with four setae. Vestigial stigmata slightly darker than sterna, Flagellum long, club shaped, expanded distally, bearing 22 major setae, Pedipalpal trochanter not produced distally; femur and patella narrow proximally, expanded distally; tibia without mesal, subapical spur. Tarsal-basitarsal spurs about

2/5, claw about 4/5 length of tarsus-basitarsus. Tarsal segments of leg I of the following approximate proportions: 23-7-9-8-8-8-26.

Other leg segment measurements given in Table 2.

Female. Flagellum with five articles.

Comparisons.--See under P_. occidental is. Distribution.--Known from various localities in Tamaulipas,

Mexico.

Additional records.--Two adult females, one juvenile, 60 mi S

Ciudad Victoria, ex bromeliads, 17 November 1948, H, B, Leech (CAS), 42

Protoschizomus occidental's Rowland, 1975

(Fig. 14; Table 2; Map 3)

Protoschizomus occidentalis Rowland, 1975:4-5.

Holotype,--An adult male, taken under a rock, 13 mi SW Colima,

Mexico, on 16 July 1972 by A, Jung (AMNH, examined),

Paratype,--A subadult female, taken with the holotype (AMNH, examined).

Description,--Male, Color brownish. Carapace with four pairs of dorsal and two apical setae, Eyespots absent, Metapeltidium divided medially into two plates. Anterior sternum with 11 entire setae. Abdominal terga I to II with two setae. III to IV with four setae, VIII to IX with six setae. Vestigial stigmata darker than sterna. Flagellum long, club shaped, expanded distally, bearing 22 major setae. Pedipalpal trochanter not produced distally; tibia without mesal, subapical spur. Tarsal-basitarsal spurs about 2/5, claw about 4/5 length of tarsus-basitarsus. Tarsal-basitarsal segments of leg I of the following approximate proportions:

24-7-8-7-8-7-27, Other leg measurements given in Table 2,

Female unknown.

Comparisons,--The two species of this genus can be separated most easily by comparison of the male's flagella. It is distally more expanded in P^, pachypalpus than in P^. occidentalis. The carapace of P_. pachypalpus has five pairs of dorsal carapacal setae whereas P^. occidentalis has four pairs. The eighth abdominal tergum 43 of the former has 10 setae, but only six exist in the latter. Other minor setational differences occur.

Distribution.--P^. occidentalis is known from near Colima,

Colima, Mexico.

Agastoschizomus lucifer Rowland, 1971

(Figs. 1, 2, 16; Table 2; Map 3)

Agastoschizomus lucifer Rowland, 1971:13-14, 15-17; Rowland, 1973a:10;

Rowland, 1973b:197, 200-201, 202; Dumitresco, 1974:282;

Brignoli, 1974:150; Rowland, 1975:9,

Holotype,--An adult male, taken in Sotano de la Tinaja, 10 km

NNE Ciudad Valles, San Luis Potosi, Mexico, on 9 April 1966, by J,

Fish and D, McKenzie (AMNH, examined).

Allotype,--An adult female, taken in Sotano de Yerbaniz,

22.5 km N Ciudad Valles, San Luis Potosi, Mexico, on 28 March 1970,

by W, Elliott (AMNH, examined),

Paratypes,--An adult female taken with the allotype (AMNH, examined); and adult male taken in Sotano de Yerbaniz, on 8 January

1971, by W, Elliott (AMNH, examined).

Description,--Male, Color brownish. Carapace with two pairs

of dorsal and three apical setae, Eyespots absent, Metapeltidium divided medially into two plates by a very narrow suture. Anterior

sternum with nine entire setae. Abdominal terga I to V with two setae,

terga VI to VII with four setae, terga VIII to IX with six setae.

Vestigial stigmata darker than sterna, Flagellum long, cylindrical. 44 apically modified, Pedipalpal trochanter not produced distally; femur and patella bearing several stout spines, tibia without mesal, subapical spur. Tarsal-basitarsal spurs about 2/3, claw about 7/6 length of tarsus-basitarsus. Tarsal-basitarsal segments of leg I of the following approximate proportions: 60-19-18-16-21-56. Other leg segment measurements given in Table 2.

Female. Flagellum with five articles.

Comparisons,--See under A^. huitzmolotitlensis.

Remarks.--See under A. huitzmolotitlensis.

Distribution.--)^, lucifer is known from caves in the southern portion of Sierra de El Abra, San Luis Potosi, Mexico.

Additional Records.--SAN LUIS POTOSI: El Sotano de Matapalma,

21 km N Ciudad Valles; one juvenile, 30 December 1972, R, Fieseler

(AMNH); one juvenile, 29 May 1969, R. Mitchell, F. Abernathy, T.

Albert (Af-INH). El Sotano de la Tinaja, 10 km NNE Ciudad Valles; two males, one female, four juveniles, 18 February 1970, J, A, L. Cooke

(AMNH); 18 February 1970, R. W. Mitchell, two females, one juvenile

(TTU); 16 March 1972, D. Kiser, one juvenile (TTU); one juvenile,

9 April 1966, J. Fish, D. McKenzie (AMNH); one female, one juvenile,

7 January 1970, R. W. Mitchell (AMNH). El Sotano de Yerbaniz, 22.5 km

N Ciudad Valles; one female, four juveniles, 7 January 1970, S. Wiley

(TTU); one male, one female, 7 January 1970, W, Elliott (TTU); one juvenile, 8 January 1970, W. Elliott (TTU); one female, one juvenile,

9 January 1970, W, Elliott (TTU); one female, nine juveniles, 17 45

February 1970, R, W. Mitchell (TTU); four juveniles, 28 March 1970,

W. Elliott (TTU),

Agastoschizomus huitzmolotitlensis Rowland, 1975

(Fig, 17; Table 2; Map 3)

Agastoschizomus huitzmolotitlensis Rowland, 1975:8-10,

Holotype,--An adult male, taken in Sotano de Huitzmolotitla,

2 km SW Tlamaya, and approximately 10 km N Xilitla, San Luis Potosi,

Mexico, on 24 January 1964, by Terry Raines and Tommy Phillips, in mud room 9000 feet from entrance (AMNH, examined).

Description,--Male. Color brownish. Carapace with one pair of dorsal and three apical setae, Eyespots absent, Metapeltidium undivided, slightly emarginate posteriorly. Anterior sternum with

11 entire setae. Abdominal terga I to IV with two setae, terga V to VII with four setae, terga VIII and IX with 10 and six setae, respectively. Vestigial stigmata slightly darker than sterna,

Flagellum long, cylindrical, bearing 18 major setae, apically modified,

Pedipalpal trochanter not produced distally; femur and patella bearing

several stout spines; tibia without mesal, subapical spur,

Tarsal-basitarsal spurs about 2/3, claw about 7/6 length of

tarsus-basitarsus, Tarsal-basitarsal segments of Leg I of the

following approximate proportions: 45-13-14-14-15-13-43, Other leg measurements given in Table 2,

Female unknown. Figs. 14-19,--Parts of protoschizomids: 14-17, lateral views of male's flagella: 14, P^, occidentalis; 15, P^, pachypalpus; 16, A^, lucifer; 17, A^, huitzmolotitlensis; 18, lateral view of right pedipalp of female P^, pachypalpus; 19, lateral view of female's flagellum of P^, pachypalpus. Scale: Figs, 14-17, 19, 1 cm = ,12 mm;

Fig, 18, 1 cm = .25 nrn. 47 Table 2.--Measurements of the members of the Protoschizomidae:

1, one male, P^. pachypalpus; 2, one female, P^, pachypalpus; 3, one male, P^, occidentalis; 4, four males, A^, lucifer; 5, four females,

A, lucifer; 6, one male. A, huitzmolotitlensis. 49

TABLE 2 Taxa

n 1 2 3 4 5 6 1 1,24 1,16 1,16 2.29-2,41 2,84-3,21 1.84 2 ,61 ,46 ,53 1.37-1,40 1,26-1,43 .98 3 ,28 - ,25 ,41-,42 - .26 4 ,97 ,79 1,02 3.36-3,51 3,48-3,76 2.53 5 .80 ,71 ,80 3,99-4.14 3,80-4,16 3.11 6 ,71 ,69 .74 3,24-3,38 2,90-3,30 2,50 7 1.05 ,80 ,99 1.97-2.10 2,04-2,16 1.54 8 1,06 ,70 1,07 2,46-2,66 2,46-3.01 1,83 9 .44 .39 ,48 1.32-1.40 1.38-1.56 1.05 10 ,34 ,34 ,38 1.69-1.86 1,75-1,97 1,35

11 ,52 ,44 ,54 1.18-1,30 1.25-1.33 .87

12 ,95 ,80 .98 2.41-2,62 2.54-2.90 1.88

13 ,53 ,44 ,48 1.13-1.20 1.11-1,43 .96

14 ,37 ,36 ,36 1,68-1.72 1,74-1,92 1,24

15 .77 .66 .70 1,41-1,45 1,51-1.54 1,04

16 .89 ,81 ,83 3,18-3.34 3,32-3.62 2,24

17 .38 ,30 ,35 1.51-1.54 1.57-1.77 1,15

18 .43 ,35 .41 2.61-2.68 2.62-2.86 1,90

19 .63 ,49 - 1.75-1.83 1.75-2.10 1,42 50

Comparisons,--A^, huitzmolotitlensis is somewhat smaller and displays more advanced troglobite facies than A, lucifer. The flagellum in particular is relatively much narrower basally. The latter, while perhaps not directly related to cave adaptation, serves best to distinguish the two species of the genus.

Distribution,--This species is known only from Sotano de

Huitzmolotitla, near Xilitla, San Luis Potosi, Mexico,

Remarks,--The two species of the genus Agastoschizomus are remarkable troglobites. They are extremely large species, without parallel in the rest of the order. Their pronounced morphological similarities lead me to believe that it is highly unlikely that they are derived from an immediate common ancestor morphologically much different from either one of them. This is important in considering the origin of these troglobites. Though the caves which these two species inhabit are proximal, separated by some 70 km, they are of distinctly different geological associations. The common ancestor of A, lucifer and A^, huitzmolotitlensis must have been an epigean species. The extreme size and pedipalpal configurations common to these species and to their hypothetical immediate ancestor are probably not totally due to adaptations to cave environments.

The cubanicus group

Members of this group are characterized by moderate to large

size (,96-1,37 carapacal length). The color varies from brown to Table 3,—Comparisons of the members of the cubanicus group.

For explanation of the characters see Phylogeny, 52

O) to C 3 o .a r— o (A E

+JT3

r- (0 c-o •as 1- 0)

•p +*

1- C O 3 -O •r- in o u 0) C o p r- (Q

tA p

V) U > 0) -3 5-

0) c •— to o) a. C 3 s r- O •r- 0)

r— O

lA M Z3 c O 0) u

1

«CUJ < Z LU Sz z* O UJ t;3 a c/1 a. u. Maps 1-2,—Distribution of schizomids of the cubanicus and simonis groups: Map i , cubanicus group members: 1, S^. cubanicoides;

2, S^. cubanicus; 3, S^, biconourus; 4, S^, subiconourus; 5, S^, cousinensis; 6, S^, primibiconourus; 7, S_, peckorum; 8, S^. viridis;

9, S^, brevipatellatus; 10, S^, longipalpus; 11, S^, desecheo; 12,

S^. monensis; 13, S^, insignis. Map 2, simonis group members: 1,

S_, mumai; 2, S_, central is; 3, S^, simonis; 4, S^, flavescens; 5,

S_, dispar; 6, S^. tobago; 7, S_, trinidanus; 8, S^. drakos; 9, OTU #1,

OTU #2; 10, S^, gladiator; 11, S_, vanderdrifti, S^, surinamensis.

Question marks indicate species doubtfully placed in the simonis group. 54

X 1

V <" c^^^^i^^'"'•

^ •

< .>^ ;>'*»*

'.^.^'^X -o.t'. J-.-/ /

71 55 green. The eyespots vary from irregular to triangular and indistinct to well defined. The carapace may have two or three pairs of dorsal and two apical setae. The arrangement of the setae may be variable and the middle pair may be smaller than the others. The abdomen shows no elongation of posterior segments. The pygydial segments are yery shallow. The males of most species have a well developed posterodorsal abdominal process, which is usually truncate, but may vary within the species from truncate to round. The flagellum, with two exceptions bears a central to subapical median depression.

Several species have bilateral projections arising from above and beside this depression. In some species the flagellum is robust and globose, but is usually small. The female's flagellum is moderate to long (.19-.36), and composed of four articles. The female's spermathecae are characterized by elongation of the laterals and reduction of the medians. In a few species the lateral spermathecae are very large and terminate in a spherical, sclerotized bulb. In other species the median spermathecae are reduced. The pedipalps are always sexually dimorphic. Males of certain species have highly variable pedipalps. The trochanter is usually in some way elaborated, and the femur, patella, and tibia are usually elongate, but the latter has no spur apposible to the tarsus-basitarsus.

Distribution,--Central America: Costa Rica, Antilles: Cuba,

Jamaica, Haiti, Puerto Rico, Martinique, 56

Remarks.--A species which probably belongs in this group, but which I have not seen, is S^, negreai Dumitresco, 1973, On the basis of her description this species appears to be most closely related to

S^, biconourus, another Cuban species. I have also been unable to examine S^. dispar Hansen and Sorensen, 1905, which may also belong in this group. It is known by females only and details of its anatomy are inadequately described. It was taken in Martinique and may be a close relative of S^. insignis.

Subordinate taxa.--Cubanicus complex: S^, cubanicus, S^. cubanicoides; primibiconourus complex: S^. cousinensis, S^. primibiconourus, S^. longipalpus, S^. brevipatellatus; viridis complex:

S^. subiconourus, S^. monensis, S^. desecheo, S^, biconourus, S^, insignis,

S^, peckorum, S^, viridis,

Schizomus cubanicus, new species

(Figs, 21, 37, 58; Tables 3, 4; Map 1)

Holotype.--An adult male, taken at San Antonio de los Banos,

Habana, Cuba, on 4 June 1972, by Luis F. de Armas (IZACC).

Description.--Male, Color brownish. Carapace with two pairs of dorsal and two apical setae, Eyespots indistinct, oblong with irregular margins. Anterior sternum with 13 entire setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segment

XII with slight posterodorsal process. Vestigial stigmata slightly darker than sterna, Flagellum lanceolate, dorsally compressed, with a single median depression subapically. Pedipalpal trochanter much 57 elongate, not produced distally, femur, patella, and tibia all elongate, no segments with additional armature, though single enlarged setae apposes the tarsus-basitarsus, Tarsal-basitarsal

spurs about 1/10, claw about 1/4 length of tarsus-basitarsus.

Tarsal-basitarsal segments of leg I of the following approximate proportions: 39-6-8-7-8-9-22, Other leg segment measurements given

in Table 4.

Female unknown.

Comparisons.--S^. cubanicus is larger and has proportionately

longer pedipalps and first legs than S^. cubanicoides, its closest relative. The variability in pedipalp length in other species,

however, and lack of adequate sample size of either of these species make diagnosis on the basis of pedipalp length questionable. The greater development of basolateral elevations on the flagellum of

S^, cubanicus is probably more reliable. The latter species also

shows a greater development of the posterodorsal process. S^.

cubanicoides lacks the large tibial seta on the pedipalps which

apposes the tarsus-basitarsus in S^. cubanicus.

Distribution,--This species is known only from the type

locality.

Etymology,--The specific name is an adjectival form taken

from Cuba,

Remarks,—While the female of this species is unknown, it

will probably bear distinct similarities to females of S^. cubanicoides. 58

Females of the latter species are notable in that they have four pairs of dorsal carapacal setae, while the males have two pairs as in S^. cubanicus,

Schizomus cubanicoides, new species

(Figs, 20, 38, 60; Tables 3, 4; Map 1)

Holotype.--An adult male, taken at Coto, Costa Rica, on 5

July 1957, by E. Dixon (AMNH),

Allotype,--An adult female, taken with the holotype (AMNH),

Paratypes,--An adult male, female, and two immatures, taken at Coto, Costa Rica, on 4 September 1957, by E, Dixon; an adult male taken with the holotype (AMNH); an adult female and two immatures, taken at Golfito, Costa Rica, on 27 August 1957, by E. Dixon (AMNH),

Description.--Male. Color brownish. Carapace with two pairs of dorsal and two apical setae. Eyespots indistinct, oblong, with irregular margins. Anterior sternum with 10 entire setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segment XII with slight evidence of posterodorsal process. Vestigial stigmata hardly distinct in color from sterna. Flagellum lanceolate, dorsally compressed, with a single median depression subapically. Pedipalpal trochanter much elongate, but not produced distally, femur, patella, and tibia all elongate, no segments with additional armature.

Tarsal-basitarsal spurs about 1/10, claw about 1/3 length of tarsus-basitarsus, Tarsal-basitarsal segments of leg I of the 59 following approximate proportions: 31-5-6-6-8-8-16, Other leg segment measurements given in Table 4.

Female. Carapace with four pairs of dorsal setae. Flagellum with four sections, Pedipalps not elongate. Lateral spermathecae about twice as long as medians, very long recurved, the tips with enlarged, sclerotized bulbs, medians very weak, with no areas of heavy sclerotization.

Comparisons.--See comparisons under S^. cubanicus.

Distribution,--This species is known from Coto and Golfito,

Costa Rica.

Etymology.--The specific name means like cubanicus.

Remarks,--S^. cubanicoides is unique in having a different number of dorsal carapacal setae in the sexes. The fact that males and females have been collected together on two occasions, however, supports the assumption that they actually represent sexes of the same species.

Schizomus cousinensis, new species

(Figs. 24, 41, 79; Tables 3, 4; Map 1)

Holotype.--An adult male, taken in Cousin's Cove Cave #1,

Cousin's Cove, Hanover Parish, Jamaica, on 31 August 1974, by S.

Peck (AMNH).

Allotype.--An adult female, taken with the holotype (AMNH).

Paratypes.--An adult male and female, taken with the holotype

(AMNH), 60

Description,--Male, Color brownish. Carapace with two pairs of dorsal and two apical setae, Eyespots indistinct. Anterior sternum with 11 bifid setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segment XII with slightly rounded posterodorsal process. Vestigial stigmata darker than sterna,

Flagellum spade shaped, with a pair of subdistal depressions flanked by two small, but well defined elevations, Pedipalpal trochanter not produced distally, but it and other palpal segments elongate, but without armature, Tarsal-basitarsal spurs about 1/6, claw about 1/4 length of tarsus-basitarsus. Tarsal-basitarsal segments of leg I of the following approximate proportions: 35-7-8-6-8-8-18.

Other leg measurements given in Table 4.

Female. Carapace with three pairs of dorsal setae, Flagellum with four sections, Pedipalps not elongate. Lateral spermathecae twice length of medians. Laterals terminate in large, highly sclerotized bulbs.

Comparisons,--See comparisons under S^. primibiconourus.

Distribution,--This species is known only from Cousin's Cove

Cave #1, Hanover Parish, Jamaica,

Etymology.--The specific name is an adjectival form taken from Cousin's Cove Cave, meaning belonging to.

Remarks,--See remarks under S^, primibiconourus.

Variations,--The holotype possesses extremely long pedipalps, but those of the male paratype are short and yery much like those of the allotype and female paratype. 61

Schizomus primibiconourus, new species

(Figs. 26, 42, 50, 79; Tables 3, 4; Map 1)

Holotype,--An adult male, taken in Oxford Cave, Auchtembeddie,

Manchester Parish, Jamaica, 4-8 April 1968, by S. Peck and A. Fiske

(AMNH),

Paratype.--An adult male, taken with the holotype (AMNH); an adult male taken in St, Claire Cave, 1,5 mi SW Ewarton, St, Catherine

Parish, Jamaica, on 7 April 1968, by S. Peck and A. Fiske (AMNH),

Description,--Male. Color brownish. Carapace with two pairs of dorsal and two apical setae. Eyespots indistinct. Anterior sternum with 11 bifid setae. Abdominal terga I-VII with two setae, terga

VIII-IX with four setae, segment XII with slight emargination of posterodorsal process. Vestigial stigmata darker than sterna,

Flagellum spade shaped, with distal depression flanked by two small, but well developed elevations, Pedipalpal trochanter slightly produced apically, not elongate, other segments slightly elongate,

but without armature, Tarsal-basitarsal spurs about 1/5, claw about

1/3 length of tarsus-basitarsus, Tarsal-basitarsal segments of leg I of the following approximate proportions: 43-7-8-11-9-10-19. Other

leg segment measurements given in Table 4.

Female unknown.

Comparisons.--This species is most closely related to S^.

cousuiensXs, from which it can be distinguished by its having a

single flagellar depression and three pairs of dorsal setae in the 62 male, whereas S^. cousinensis has a pair of depressions and two pairs of dorsal setae. The latter two species are generally similar to

S^. cubanicus and S^. cubanicoides in size and development of the male's flagellum. S^. primibiconourus and the latter species have well developed subapical median depressions, but S^. primibiconourus is distinct in the development of the elevations lateral to the depressions.

Distribution,--This species is known from Oxford Cave,

Manchester Parish, and St. Claire Cave, St. Catherine Parish, Jamaica,

Etymology,--The specific name is from the Latin primi- meaning first, bX meaning two, the Greek cono- meaning cone and urus meaning tail. This name describes the primitive condition of the pair of cone shaped elevations on the flagellum of the male of this species.

Remarks,--While this species and S^. cousinensis are cladistically closer to S_, longipalpus and S_. brevipatellatus they more strongly resemble members of the cubanicus complex. This is accounted for by the remarkable variation in size and pedipalpal characters in the former species.

The middle pair of dorsal carapacal setae are apparently in a transitional state in S^, primibiconourus, but are absent in S^, cousinensis. Most species in the cubanicus group show a reduction in size of this pair. 63

Variation.--The holotype has a very small seta present on the right side where the middle pair of dorsal carapacal setae occur in other species. A paratype has one on the left side.

Schizomus longipalpus, new species

(Figs. 23, 40, 68; Tables 3, 4; Map 1)

Holotype.--An adult male, taken at La Vestite, Haiti, on

23 September 1916, by Darlington, elevation 6-7000 ft, (MCZ),

Paratypes,--Two adult males, taken at Furey, Haiti, during

1912-1913, by W, M, Mann (MCZ).

Description.--Male, Color brownish. Carapace with two pairs of dorsal and two apical setae. Eyespots vaguely triangular,

irregular, but distinct borders. Anterior sternum with 13 bifid setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segment XII with slight development of posterodorsal

process. Vestigial stigmata darker than sterna. Flagellum spade

shaped, with distomedian depression flanked basolaterally by distinct

elevations. Pedipalpal trochanter extremely long and extremely

produced distally. Other segments elongate, but unarmed, tibia with

a large setae apposible to tarsus-basitarsus. Tarsal-basitarsal

spurs about 1/10, claw about 1/4 length of tarsus-basitarsus. Tarsal

basitarsal segments of leg I of the following approximate proportions:

59-10-13-12-13-13-24, Other leg segment measurements given in Table 4.

Female unknown. 64

Comparisons,--S^, longipalpus is similar in its development of the male's pedipalps to other species of the group, except in the extreme elongation of the trochanter, both proximally and distally.

Its closest relatives are ^, primibiconourus and S^, brevipatellatus,

S^. longipalpus is much larger than S^, primibiconourus, which also has onl'y slightly elongated palpal segments and a better developed median flagellar pit, but smaller lateral elevations, S^, brevipatellatus is somewhat larger than S^. longipalpus and the flagellum lacks the median pit. The extreme elongation of the pedipalps in the latter two species are probably not homologous developments. The trochanter of S^. longipalpus is extremely long, the other segments not being out of the ordinary for the group. The elongate pedipalps of S^. brevipatellatus are achieved by attenuation of the femur and tibia, the patella being extremely short.

Remarks,--As in S^, brevipatellatus the remarkable development of the pedipalps may be related to some unusual food source. The great extension of the trochanter in effect sets the apposible parts of the pedipalps some distance from the body proper.

Distribution,--This species is known from La Vestite and

Furey, Haiti.

Etymology.--The specific name comes from the Latin longi- meaning long, a palp. 65

Schizomus brevipatellatus, new species

(Figs. 22, 39, 64, 65; Tables 3, 4; Map 1)

Holotype.--An adult male taken at Cape Haitien, Haiti, in

January 1913, by W, M, Mann (MCZ),

Description,--Male, Color brownish. Carapace with two pairs of dorsal and two apical setae. Eyespots present, but indistinct.

Anterior sternum with 13 bifid setae. Abdominal tergum I with two setae, II with four setae, III-VI with two setae, terga VII-IX with six setae, segment XII with slight development of posterodorsal process. Vestigial stigmata much darker than sterna. Flagellum lanceolate, slightly dorsally compressed, with a median swelling flanked by lateral pits. Pedipalpal trochanter slightly produced distally, but short, patella short, other segments extremely long, but unarmed. Tarsal-basitarsal spurs about 1/10, claw about 1/5 length of tarsus-basitarsus. Tarsal-basitarsal segments of leg I of the following approximate proportions: 55-9-11-11-14-14-30.

Other leg measurements given in Table 4.

Female unknown.

Comparisons.--See comparisons under S^. longipalpus.

Distribution.--This species is known only from the type locality.

Etymology.--The specific name comes from the Latin brevi- meaning short, and patella. 66

Remarks.--The form of the pedipalps is unique among the schizomids. The patella is extremely shortened, and apparently the joint connecting it with the tibia is inflexible. This unique structure probably reflects a novel function of the pedipalp, at least in so far as the apposition of articles. It may be that these limbs serve to allow the males of these species to utilize a food source which is necessarily held a good distance from the body, such as ants or other relatively defensive arthropods.

The flagellum of S^. brevipatellatus departs greatly from those of the remainder of the group. While this species is similar in other respects to S^. longipalpus I further think that the highly modified pedipalps occurred independently in the two species, since the elongation is accomplished in radically different ways in the two species.

Schizomus subiconourus, new species

(Figs. 26, 43, 51, 62, 77; Tables 3, 4; Map 1)

Holotype.--An adult male, taken at El Marra, Santiago, Cuba, on 22 May 1972, by Luis F. de Armas (IZACC).

Allotype.--An adult female taken with the holotype (IZACC).

Paratypes.--An adult male, adult female, and immature, taken with the holotype (IZACC).

Description.--Male. Color greenish. Carapace with three pairs

of evenly spaced dorsal and two apical setae. Eyespots round to distinctly triangular, well defined. Anterior sternum with 12 bifid 67 setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segment XII with rounded posterodorsal process. Vestigial stigmata slightly darker than sterna. Flagellum triangular, with slight apical depression flanked basolaterally and distally by elevations. Pedipalpal trochanter greatly enlarged and produced distally, femur elongate and produced beyond insertion of patella, patella, tibia, and tarsus-basitarsus elongate. Tarsal-basitarsal spurs about 1/8, claw about 1/3 length of tarsus-basitarsus.

Tarsal-basitarsal segments of leg I of the following approximate proportions: 30-6-6-6-7-8-19. Other leg segment measurements given in Table 4.

Female. No posterodorsal abdominal process. Flagellum with four articles. Pedipalps not elongate. Lateral spermathecae about twice as long as medians, both with terminal sclerotized bulbs.

Comparisons.--This species is most similar in several respects to S^. desecheo and S^, monensis. The latter species possess only a vague apical depression on the male's flagellum; this depression is also poorly, but better, developed in S^, subiconourus. The latter species is unique in the degree of development of the femur of the male's pedipalp which is produced beyond its insertion with the patella. In Cuba S^. subiconourus is distinguished from other males by the reduction of the median apical depression which is well developed in S^. cubanicus and S^. biconourus. Males of S^. rowlandi apparently lack a median pit, but also lack dorsal elevations, as is the case in S. armasi. 68

Distribution.--This species is known only from the type locality.

Etymology.--The specific name is from the Latin sub meaning under, bi_ meaning two, the Greek cono- meaning cone and urus meaning tail. This name describes the underdeveloped condition of the pair of cone shaped elevations on the flagellum of the male of this species.

Remarks,--The articulation of the femur-patellar joint of the pedipalp in male schizomids is probably usually flexible through nearly 180°, The modification of this joint is such that flexure much past a right angle is impossible, S^, longipalpus and S^, brevipatellatus also have unique development of the male's pedipalp.

Variation.--The eyespots of this species may vary from round to triangular. So few specimens are available, however, that the typical condition, if one exists, cannot be ascertained.

Schizomus monensis, new species

(Figs, 28, 45, 78; Tables 3, 4; Map 1)

Holotype,--An adult male, taken in Cueva del Agua, Mona

Island, Puerto Rico, on 3 June 1974, by S, Peck (AMNH),

Allotype.--An adult female, taken with the holotype (AMNH).

Paratypes.--An adult male, four adult females, and an immature taken in Cueva Dona Gena, Mona Island, on 5 June 1974, by S, Peck

(AMNH); an adult female taken in Cueva Negro, Mona Island, on 3 June

1974, by S. Peck (AMNH). 69

Description,--Male. Color greenish. Carapace with three pairs of dorsal and two apical setae, Eyespots indistinct. Anterior sternum with 11 bifid setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segment XII with well developed, truncate posterodorsal process. Vestigial stigmata darker than sterna,

Flagellum spade shaped, thick, with two mediolateral and distal vague elevations, median pit vague or absent, Pedipalpal trochanter produced distally, no segments elongate, Tarsal-basitarsal spurs about 1/6, claw about 1/3 length of tarsus-basitarsus, Tarsal-basitarsal segments of leg I of the following approximate proportions: 37-6-7-7-8-

10-19. Other leg segment measurements given in Table 4.

Female. Flagellum with four articles. Lateral spermathecae about one and one half times as long as medians, laterals terminate in large sclerotized bulbs.

Comparisons,--Males of this species can be told from those of its closest relative, S^. desecheo, by the much thicker flagellum and the less distinct eyespots. The middle pair of dorsal carapacal setae are smaller in S^, monensis. Otherwise the species are very similar.

Also, see comparisons under S^, subiconourus.

Distribution,--This species is known from three caves on Mona

Island, Puerto Rico,

Etymology,--The specific name is an adjectival form taken from Mona Island, meaning belonging to.

Variation.--The female allotype has four pairs of dorsal carapacal setae, whereas the other female paratypes have three pairs. 70

Schizomus desecheo, new species

(Figs, 27, 44, 67; Tables 3, 4; Map 1)

Holotype.--An adult male, taken on Desecheo Island, Puerto

Rico, on 29 March 1961, by J. R. Gorham (MCZ).

Paratype.--An immature, taken with the holotype (MCZ).

Description.--Male. Color greenish. Carapace with three pairs of dorsal and two apical setae. Eyespots distinct, angular. Anterior sternum with 10 bifid setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segment XII with well developed, truncate posterodorsal process. Vestigial stigmata distinctly darker than sterna, Flagellum lanceolate, somewhat dorsally compressed with two mediolateral distinct elevations, median pit vague or absent,

Pedipalpal trochanter slightly elongate, not produced distally, all other segments only slightly elongate, Tarsal-basitarsal spurs about

1/7, claw about 1/3 length of tarsus-basitarsus. Tarsal-basitarsal segments of leg I of the following approximate proportions: 38-8-8-9-

9-10-23. Other leg segment measurements given in Table 4,

Female unknown.

Comparisons,--See comparisons under S^, monensis and S^,

subiconourus.

Distribution,--This species is known from a single collection from Desecheo Island, Puerto Rico,

Etymology,--The specific name is an adjectival form taken

from Desecheo Island, meaning belonging to. 71

Schizomus biconourus, new species

(Figs. 34, 52, 53, 61, 66, 76; Tables 3, 4; Map 1)

Holotype,--An adult male, taken at Cienfuegos, Soledad, Cuba, on 3 September 1972, by Luis F, de Armas (IZACC),

Allotype,--An adult female, taken with the holotype (IZACC),

Paratype.--An adult male, taken in Soledad, Cuba, on 31 July

1933, by J. A. Weber (MCZ).

Description.--Male. Color greenish. Carapace with three pairs of dorsal and two apical setae, the dorsals in a tight basal group,

Eyespots ovoid. Anterior sternum with 12 bifid setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segment XII with well developed, truncate posterodorsal process. Vestigial stigmata lighter than sterna, Flagellum triangular, with distal depression flanked by pair of well developed elevations. Pedipalpal trochanter produced distally, armed with a dorsal spur, other segments elongate, but unarmed, Tarsal-basitarsal spurs about 1/8, claw about

1/3 length of tarsus-basitarsus, Tarsal-basitarsal segments of leg I of the following approximate proportions: 34-6-7-8-7-10-21, Other

leg segment measurements given in Table 4.

Female. Dorsal carapacal setae not grouped tightly.

Flagellum with four articles. Pedipalps not elongate or armed.

Lateral spermathecae somewhat longer than the medians, the origins

closely situated and lobes slightly overlapping in ventral view.

Apex of lateral and median spermathecae expanded into slightly

sclerotized bulbs. 72

Comparisons,—S^, biconourus has a great development of the elevations proximal to the distal flagellar depressions in the males.

Another Cuban species, S^, negreai Dumitresco, 1973, which I have not seen, seems to possess a lesser, but nonetheless distinct development of these elevations. Illustrations of the latter species show the flagellum to be much less triangular in shape than S^, biconourus.

The spermathecae of the two species are apparently very similar.

Distribution.--This species is known from Soledad, Cuba.

Etymology.--The specific name is from the Latin bi meaning two, the Greek cono- meaning cone and urus meaning tail. This name describes the morphology of the flagellum of the male of this species.

Schizomus insignis Hansen and Sorensen, 1905

(Figs. 36, 47, 54, 59, 82; Tables 3, 4; Map 1)

Schizomus insignis Hansen and Sorensen, 1905:11, 26, 39, 47-49, 50,

74; Mello-Leitao, 1931:18; Giltay. 1935:6; Takashima, 1943:93;

Remy, 1961:506,

Cotypes,--An adult male and immature, taken in Martinique, by

E, Simon (UZM, examined); an adult female, taken in Martinique, by E,

Simon (MNHN, examined).

Description.--Male. Color greenish. Carapace with three pairs of dorsal and two apical setae. Eyespots oval, but indistinct.

Anterior sternum with 11 entire setae. Abdominal terga I-VII with

two setae, terga VIII-IX with four setae, segment XII with well developed, truncate posterodorsal process. Vestigial stigmata almost 73 indistinguishable from sterna, Flagellum spade shaped, with two lateral and a median process separated by a median pit, Pedipalpal trochanter greatly elongate, but not produced distally, other segments elongate and unarmed, Tarsal-basitarsal spurs about 1/10, claw about 1/6 length of tarsus-basitarsus, Tarsal-basitarsal segments of leg I of the following proportions: 36-7-8-8-7-11-18,

Other leg segment measurements given in Table 4.

Female, Flagellum with four articles, Pedipalps not elongate. Lateral spermathecae about four times longer than medians.

Laterals terminate in highly sclerotized bulbs,

Comparisons,--S^. insignis is about the size and has similar pedipalps to those of S^. cubanicoides and S^, cubanicus. The latter species are more primitive, however, and lack the greenish color, the pedipalpal trochanter spur, and the median pair of dorsal setae.

Distribution,--This species is known from Martinique, Lesser

Antilles.

Remarks.--The eyespots are rather hard to see since the types are now quite bleached. Hansen and Sorensen (1905) describe them as

"moderately large, short but broad, whitish." How they compare to other species of the cubanicus group is uncertain. 74

Schizomus peckorum, new species

(Figs. 35, 48, 55, 73, 80, 81; Tables 3, 4; Map 1)

Holotype.--An adult male, taken in Windsor Great Cave,

Windsor, 10 mi. S Falmouth, Trelawney Parish, Jamaica, on 5 April

1968 by S. Peck and A. Fiske (AMNH).

Allotype.--An adult female, taken with the holotype (AMNH).

Paratypes.--An adult female, taken with the holotype (AMNH); an adult male and five adult females, taken in Mosley Hall Cave, near Guys Hill, St. Ann Parish, Jamaica, on 27 December 1972, by

S. and J. Peck.

Description.--Male. Color greenish. Carapace with three pairs of dorsal and two apical setae, the median dorsals very reduced.

Eyespots oval, well defined. Anterior sternum with 13 bifid setae.

Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segment XII with well developed, truncate posterodorsal process.

Vestigial stigmata darker than sterna. Flagellum nearly globose, with two lateral and one median processes separated by median pit.

Pedipalpal trochanter produced distally, other segments slightly

elongate, but unarmed. Tarsal-basitarsal spurs about 1/7, claw about

1/3 length of tarsus-basitarsus. Tarsal-basitarsal segments of leg I

of the following approximate proportions: 50-8-11-10-11-12-23. Other

leg segment measurements given in Table 4.

Female. Flagellum with four articles. Pedipalps not

elongate. Lateral spermathecae about five times length of medians. 75 laterals with extreme development of sclerotized bulbs.

Comparisons.--S^. peckorum is most similar to S^. viridis in several respects. S^. peckorum males have a smaller flagellum, with better defined elevations. The female's spermathecae of S^. peckorum has much larger sclerotized terminal bulbs. In S^. viridis the laterals have small bulbs and the medians are only slightly shorter.

Distribution.--This species is known from caves in Trelawney and St. Ann Parishes, Jamaica,

Etymology,--The specific name is a patronym given for Stewart and James Peck, collectors of this species,

Schizomus viridis, new species

(Figs, 29-33, 49, 56, 57, 69-72, 83-86; Tables 3, 4; Map 1)

Holotype.--An adult male, taken in Abbey Cave, 2,5 mi. SW

Mandeville, Manchester Parish, Jamaica, on 24 December 1973, by

S. and J. Peck (AMNH),

Allotype,--An adult female, taken with the holotype (AMNH),

Paratypes,--An adult male, three adult females, and six

immatures, taken with the holotype (AMNH),

Description,--Male. Color green. Carapace with three pairs of dorsal and two apical setae, the median dorsal pair extremely

reduced. Eyespots triangular and well defined. Anterior sternum

with 13 bifid setae. Abdominal terga I-VII with two setae, terga

VIII-IX with four setae, segment XII with well developed, round to

truncate posterodorsal process. Vestigial stigmata darker than 76 sterna. Flagellum globose, with two lateral and a median poorly defined process, median pit present, but vague. Pedipalpal trochanter produced distally, the other segments elongate. Tarsal-basitarsal spurs about 1/7, claw about 1/3 length of tarsus-basitarsus,

Tarsal-basitarsal segments of leg I of the following approximate proportions: 52-8-11-11-10-12-23. Other leg segment measurements given in Table 4,

Female, Flagellum with four articles. Pedipalps not elongate. Lateral spermathecae two or three times longer than medians, the apex of the medians and laterals with small sclerotized bulbs.

Comparisons.--See comparisons under S^. peckorum.

Distribution,--This species is known from St, Ann, St,

Catherine, St, Claire, Clarendon, St, Elizabeth, Manchester, and

Trelawney Parishes, Jamaica.

Etymology,--The specific name is from the Latin viridi- meaning green.

Remarks,--This species is the most highly advanced cubanicus group member and the widest spread species in Jamaica, It inhabits both caves and epigean habitats and has been cullected with members of S^, portoricensis, S^, primibiconourus, and S. troglobius. As mentioned above, this species may actually represent several geographical isolates.

Variation.--There seems to be substantial geographic variation in the male's flagellum and pedipalps ,ind the female's 77 spermathecae of this species.

Additional records,--St, Ann Parish: one adult male,

Chesterfield Cave, 27 March 1973, R. Norton, R, Zimmerman (TTU); three adult males, one adult female, five immatures. Ken Connell

Hole, 8 mi, S Claremont, 19 August 1974, S, Peck (MCZ); one adult male, one immature male, Douglas Castle, Brambribo Cave, 18 August

1974, S, Peck (AMNH); one adult male, two immatures, Douglas Castle,

Falling Cave, 18 August 1974, S, Peck (TTU); three adult males, one adult female, five immatures, Hutchinson Hole Cave, 27 March 1973,

R. Norton, R. Zimmerman (AMNH); seven adult males, six adult females, five immatures, Thatchfield Light Hole, 28 March 1973, R, Norton,

R, Zimmerman (TTU); two females, Thatchfield Great Cave, 24 October

1973, R. Norton (AMNH); four males, two females, 12 immatures. Cave

River Cave, Aenon Town, 2 September 1974, S, Peck (TTU); one male, four females, four immatures, Mt, Plenty Cave, Goshen, 20 August

1974, S, Peck (AMNH); one female, Goshen, 1500 ft,, 25 December 1973,

S. and J, Peck (AMNH), St, Catherine Parish: three males, one female, two immatures, Swansea Cave, 4 November 1973, R. Norton (TTU); one male, seven females, five immatures, St, Claire Cave, 1,5 mi, SW

Ewarton, 7 April 1968, S, Peck, A, Fiske (TTU). Clarendon Parish: one female, one immature, Jackson Bay Cave, 15 August 1974, S. Peck

(AMNH); one male, one female, one immature, Jackson Bay Cave, Jackson

Bay, 2 August 1974, S. Peck (AMNH); four males, three females, two immatures, Pedro Great Cave, Pedro River, 17 August 1974, S. Peck

(MCZ); three males, one female, one immature, Pedro Great Cave, Pedro Figs. 20-28,.—Dorsal views of male's flagella of the cubanicus group: 20, Schizomus cubanicoides; 21, S^. cubanicus; 22, S^.

brevipatellatus; 23, S^. longipalpus; 24, S^. cousinensis; 25, S_. primibiconourus; 26, S^. subiconourus; 27, S^. desecheo; 28, S^. monensis. Scale: 1 cm = .06 mm.

Figs, 29-33,—Dorsal views of male's flagella of S^, viridis from various localities; 29, the type locality; 30, Pedro Great Cave,

Clarendon Parish; 31, Jackson Bay Cave, Clarendon Parish; 32, Cave

River Cave, St. Ann Parish; 33, Thatchfield Light Hole, St, Ann

Parish, Scale: 1 cm = ,08 mm.

Figs. 34-44,—Male's flagella of the cubanicus group: 34-36, dorsal views: 34, S^, biconourus; 35, S^, peckorum; 36, S^, insignis;

37-44, lateral views: 37, S^. cubanicus; 38, S^, cubanicoides; 39,

S^, brevipatellatus; 40, S^, longipalpus; 41, S^. cousinensis; 42, S^, primibiconourus; 43, S^, subiconourus; 44, S^, desecheo. Scale:

Figs, 34-38, 40-44, 1 cm = ,06 mm; Fig, 39, 1 cm = ,08 mm. 83 Figs. 45-57.--Parts of male schizomids of the cubanicus group:

45-49, lateral views of flagella: 45, S^, monensis; 46, S^. biconourus;

47, S^. insignis; 48, S^, peckorum; 49, S^, viridis; 50-57, dorsal views of posterodorsal abdominal process: 50, S^. primibiconourus; 51,

S^. subiconourus; 52, 53, S^, biconourus; 54, S^, insignis; 55, S^, peckorum;

56, 57, S. viridis. Scale: 1 cm = .06 mm.

Figs, 58-60,--Lateral views of male's right pedipalps of the cubanicus group: 58, S^. cubanicus; 59, S^, insignis; 60, S^, cubanicoides. Scale: 1 cm = .06 mm. 87

^K'^K

>v-r<<^ Figs. 61-68.--Lateral views of male's right pedipalp of the cubanicus group: 61, S^, biconourus; 62, S.. subiconourus; 63, S_, primibiconourus, trochanter and femur omitted; 64, S^. brevipatellatus;

65, S^, brevipatellatus, patella only; 66, S^. biconourus; 67, S^. desecheo; 68, S^. longipalpus. Scale: -igs, 61-63, 65-67, 1 cm =

,06 mm; Figs. 64, 68, 1 cm = .12 mm. //A

-\—r\ Figs. 69-73.—Male's right pedipalps of the cubanicus group:

69-71, lateral views of S^, viridis; 72, mesal view of S^. viridis;

73, lateral view of S^, peckorum. Scale: 1 cm = .06 mm. 91 Figs. 74-86,--Female's spermathecae of the cubanicus group:

74, 75, S^. cubanicoides; 76, S^. biconourus; 77, S^, subiconourus;

78, S^, monensis; 79, S^. cousinensis; 80, 81, S^, peckorum; 82,

S^. insignis; 83-86, S^. viridis from various localities: 83, Pedro

Great Cave, Clerendon Parish; 84, the type locality; 85, Abby Cave,

Manchester Parish; 86, St, Claire Cave, St. Catherine Parish, Scale:

1 cm = .04 mm.

Table 4,--Measurements of the members of the cubanicus group:

1, one male, S^, cubanicus; 2, three males, S^, cubanicoides; 3, two fema 1 es, S^, cubanicoides; 4, two ma 1 es, S^. cousinensis; 5, two females, S^, cousinensis; 6, three males, S^, primibiconourus; 7, three males, S^. longipalpus; 8, one male, S^, brevipatellatus; 9, two males, S^, subiconourus; 10, two females, S^, subiconourus; 11, one male, S^. monensis; 12, two females, S^. monensis; 13, one male,

S^. desecheo; 14, two males, S^. biconourus; 15, one female, S^. biconourus; 16, one male, S^, insignis; 17, one female, S_, insignis;

18, two males, S^, peckorum; 19, three females, S^, peckorum; 20, three males, S^. viridis; 21, three females, S, viridis. 95

TABLE 4 Taxa n 1 2 3 4 5 6 1 1,10 .93-.98 .95-,98 .92-1.18 1.06-1.08 1,01-1.07 2 1.13 .33-.34 ,30 .36-.37 .30-.30 .33-.34

3 ,20 .17-.19 - .19-.22 - .22-.26 4 1,23 .79-.89 ,77-,82 .97-1.26 .90-.94 1.01-1,04 5 1.49 .59-.68 ,61-,62 1 .16-1.56 1,11-1,13 1,28-1,38 6 1.10 .54-.58 ,51-58 .84-1,10 ,80-82 .98-1,03 7 ,82 .82-,89 .82-,86 ,77-,90 ,71-,72 ,80-,83 8 .76 ,98-1.09 ,87-,93 .68-,85 .66-.68 ,69-.72

9 .43 .34-,39 .35-.36 ,40-,52 ,39-,40 ,36-,38 10 .50 ,25-,27 ,25-,26 ,43-,53 ,39-,42 ,39-,40 11 .45 .37-.44 .39-.41 .38-.50 .38-.39 .41-.47

12 .65 .69-,80 .65-.69 .38-.74 .57-.60 .58-.65

13 ,30 ,33-,39 .35-.36 ,25-,35 .25-,28 .30-.38

14 .34 ,27-,30 .25-.26 ,30-,39 ,29-,31 .31-.33

15 .39 ,54-.60 .55-.59 .36-,53 ,36-,39 .39-.46

16 1.05 ,65-,68 .64-.64 .91-1,11 ,91-,97 1.00-1.06

17 .48 ,32-.38 .35-.36 ,42-.56 ,43-,45 .39-.44

18 .76 .32-.39 .36-.36 ,61-,74 ,60-,64 .62-.69

19 .65 .50-,54 .50-.52 ,54-.71 ,54-,54 .56-.62 96

TABLE 4 (conti nued)

Taxa n 7 8 9 10 11 12 1 1.24-1.35 1,36 .92-.93 .99-1,01 1.16 1,22-1,28 2 .44-,47 .55 ,30-,31 ,23-,23 .43 ,33-.33

3 ,26-,27 .27 ,23-,23 - .26 - 4 1,50-1.56 1.60 ,89-.92 ,85-,87 1.32 1.28-1.33 5 1.87-1.96 2,11 1 ,09-1.15 1,02-1,03 1.66 1.59-1.65

6 1.36-1.39 1,59 ,79-,84 ,75-.77 1.26 1,21-1,24 7 1.03-1.12 1,12 ,66-,67 ,66-,68 ,97 .90-.92

8 1.04-1.10 1,21 ,61-.65 ,62-,67 .86 ,86-,89 9 .62-.63 ,66 .33-,36 .34-.36 ,48 ,49-.51

10 .68-.72 .82 ,38-,41 .36-.38 ,55 ,56-,57

11 .61-.65 .67 .33-.37 .32-.33 ,53 ,54-,54

12 .84-.91 1.03 ,52-.53 .54-.54 ,73 ,77-,77

13 .43-.44 .48 .24-.26 .26-.26 ,35 ,36-.37

14 .46-.49 .57 .27-.28 .26-.29 ,44 .46-,47

15 .58-.63 .69 .35-.36 .35-.35 ,55 ,52-,54

16 1.27-1.39 .77 .85-.89 .87-.88 1,21 1.23-1,26

17 .65-.65 .34 .38-.40 .38-.41 ,53 ,53-.55

18 .82-.93 1.09 .55-.61 .58-.58 .81 .86-.88

19 .84-,86 .96 .49-.50 .50-.51 ,72 .77-.79 97

TABLE 4 (contin ued)

Taxa n 13 14 15 16 17 18 19

1 1.16 ,96-1,04 .96 1,01 1,00 1.24-1.31 1.22-1.28

2 .39 ,34-.35 .23 .37 .19 .46-.49 ,33-,37

3 .20 .33-.36 - .23 - .35-.37 -

4 1,10 .94-1,00 .80 1.03 .86 1.34-1.54 1,24-1,35

5 1,28 1,16-1.24 .96 1,26 1,06 1.75-1.86 1.45-1.62

6 1.01 .88-.93 ,75 ,87 ,75 1.19-1.35 1.07-1.13

7 ,82 .72-,72 ,66 ,71 .66 .97-1.09 .83-,87

8 ,77 .64-,70 ,57 ,71 .65 .93-1.07 ,89-,94

9 ,41 ,38-.40 ,33 ,40 .37 .51-.57 ,45-,51

10 ,49 .38-,43 ,33 ,44 .37 .58-.66 .52-,58

11 ,42 .36-.37 ,31 ,40 .33 .64-.68 .51-.57

12 .66 ,55-,56 ,44 .62 .55 .77-.89 .76-.82

13 .31 ,26-,27 ,23 .28 .26 .36-.42 .36-.38

14 .39 ,30-,31 ,26 ,26 .25 .43-.44 ,40-,43

15 .44 ,36-,40 .31 ,40 .37 .58-,65 .52-,55

16 1,07 ,87-,93 .82 ,96 .93 1.30-1,43 1.00-1.24

17 .47 ,39-,48 ,37 ,44 .41 1.09-1.28 .54-.56

18 .75 ,54-,61 .55 ,64 .51 .82-.97 .76-,85

19 - .62 ,50-,55 ,47 .59 .54 .75-.87 ,71-,74 98

TABLE 4 (continued)

Taxa n 20 21

1 1.31-1.35 1.38-1.42

2 ,43-.47 .35-.36

3 ,40-,44 -

4 1,25-1,38 1.20-1.24

5 1,52-1,69 1,36-1.48

6 l.lO-i.21 1.03-1.07

7 .99-1.00 .88-,93

8 .92-.93 .92-.96

9 .51-.56 .53-.55

10 .55-.56 .54-.56

11 ,51-,61 ,50-,54

12 ,78-,86 ,79-,85

13 .35-.41 .33-.39

14 .40-.46 .39-.44

15 .54-.59 .51-.52

16 1.27-1.34 1,27-1,31

17 .58-,63 .58-,62

18 .82-.87 .80-.86

19 .75-.79 .73-.75 99

River, 20 December 1972, S, and J. Peck (AMNH). St. Elizabeth

Parish: one male, three females, one immature, Wallingford Sink

Cave, Wallingford, 27 August 1974, S, Peck (TTU); Peru Cave, 5 mi.

ENE Santa Cruz, one female, 23 December 1972, S. and J, Peck (TTU).

Trelawney Parish: three males, one female, two immatures. Drip

Cave, 1.5 mi. SSE Stewart Town, 2 April 1968, S. Peck, A. Fiske

(MCZ); one immature, Windsor Great Cave, 29 March 1973, R. Norton

(AMNH).

The simonis group

Members of this group are characterized by moderate to great length (1.07-1.34 carapacal length). The color is brownish, Eyespots are present, but are usually indistinct. The carapace may have two to three pairs of dorsal and two apical setae. The male's abdomen is attenuate, the elongation may be limited to the pygydial segments, or may involve segments V-XII, The male's abdominal segment XII has a posterodorsal process, which is usually truncate, but in a few is rounded. The male's flagellum is longer in species with more elongate abdomen, but shorter in species with lesser attenuation, A pair of subproximal flagellar elevations, which are often undercut, are present in males of all species. The female's flagellum is usually long (,37), but may be very long (,61) and is composed of four articles. The female's spermathecae are characterized by the elongation of the lateral pair and a usually slight reduction of the medians. In most females the apex of the spermathecae, at least the T

Table 5,--Comparisons of the members of the simonis group.

For explanation of the characters see Phylogeny, 101

i— o f

1- c •O C f

9, «

§ 15 s •o

V) u S

UJ 5^ _i Z—1 <:r (—UJ . UJ Sid a.t- C9C9 IS tf o OO C9 UJ UJ K z a3 Si < UJ S3 b3 a. u. 102 laterals, have sclerotized bulbs. The pedipalps are not sexually dimorphic.

Distribution.--Central America: Costa Rica, Panama. South

America: Venezuela, Trinidad, Tobago, British Guiana.

Remarks.--Species which may belong to this group, but which

I have not studied include S^. gladiator Remy, 1961, S^. surinamensis

Remy, 1961, S^. vanderdrifti Remy, 1961, and S^. dispar Hansen and

Sorensen, 1905. The latter is from Martinique and all the former are from Surinam. Also see Remarks under S^. drakos.

Subordinate taxa.--Drakos complex: OTU #1, OTU #2, S_, drakos; simonis complex: S^. simonis, S^. trinidanus, S^, acrocaudatus, S^, flavescens; centralis complex: S^. tobago, S^, mumai, S^, centralis.

Schizomus sp,, OTU #1

(Fig. 106; Tables 5, 6; Map 2)

Specimen examined,--An adult female taken at Atkinson Field,

British Guiana, on 8 November 1959 (AMNH).

Description.--Male unknown. Female as follows: Color brownish. Carapace with three pairs of dorsal setae, the middle pair smallest, and two apical setae. Eyespots indistinct. Anterior sternum with 10 bifid setae, posterior sternum with bifid setae.

Abdominal terga I-VIII with two setae, tergum IX with four setae.

Vestigial stigmata darker than sterna. Flagellum missing. Pedipalpal trochanter not produced apically, Tarsal-basitarsal spur about 1/4, claw about 1/2 length of tarsus-basitarsus. Tarsal-basitarsal 103 segments of leg I of the following approximate proportions: 33-4-6-7-

7-7-17. Other leg segment measurements given in Table 6, Median and lateral spermathecae about same size; wide basally to apically; no localized sclerotization; laterals bent outward basally; medians convergent, laterals divergent.

Variation.--Only a single specimen was available for study.

Remarks,--OTU #1 is most closely related to OTU #2 with which it shares a unique development of the spermathecae. The spermathecae are short and wide basally, slightly tapering distally, the medians convergent, the lateral divergent, but of about equal size,

OTU #1 is perhaps best distinguished from OTU #2 by having no sclerotized basal portion under the spermathecae. The shapes and configuration of the spermathecae of these two species is similar.

Distribution.--This species is known only from Atkinson Field,

British Guiana.

Schizomus sp., OTU #2

(Figs. 104, 105; Tables 5, 6; Map 2)

Specimens examined.--An adult female, taken in the Bartica

District, British Guiana, on 6 May 1924 (AMNH), and two adult females, taken in Kartabo 1, British Guiana, in 1919, by A. Emerson (AMNH),

Description,--Male unknown. Female as follows: Color brownish. Carapace with three pairs of dorsal setae, the middle pair smallest, and two apical setae. Eyespots indistinct. Anterior sternum with 11 bifid setae, posterior sternum with bifid setae. 104

Abdominal terga I-VIII with two setae, tergum IX with four setae.

Vestigial stigmata darker than sterna, Flagellum with four sections.

Pedipalpal trochanter not produced distally. Tarsal-basitarsal spurs about 1/4, claw about 1/2 length of tarsus-basitarsus. Tarsal- basitarsal segments of leg I of the following approximate proportions:

35-5-8-8-8-8-17. Other leg segment measurements given in Table 6.

Median and lateral spermathecae similar in shape, but laterals longer, both pair wide basally, slightly narrower apically, ending in a small cleft; no special sclerotization; medians convergent, laterals divergent; medians and laterals basally connected to a basal piece.

Variation,--The spermathecae of the three specimens examined show some marked differences. In the specimen from the Bartica

District the medians are smaller than the laterals and the basal portion is smaller than either pair of spermathecae. The spermathecae of the Kartabo 1 specimens are larger and the medians and laterals are more nearly equal in size, although there is a marked asymmetry in the size of the laterals. The basal portion of the spermathecae is larger than either the laterals or medians.

Remarks,--It is possible that these two collections represent different, though closely related species. Without males of the species in question, or additional females from these and other localities, it is more convenient to consider tham as representing a single species. 105

Distribution.--This species is known from Bartica District and Kartabo 1, British Guiana.

Schizomus drakos, new species

(Fig. 89; Tables 5, 6; Map 2)

Holotype.--An adult male, taken in Kartabo, Bartica District,

British Guiana, on 12 October 1920 (AMNH).

Description.--Male. Color brownish. Carapace with three pairs of similar dorsal and two apical setae, Eyespots oval, indistinct.

Anterior sternum with 12 bifid setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae; abdominal segments V-XII extremely elongate; segment XII with rounded posterodorsal process.

Vestigial stigmata darker than sterna. Flagellum nearly triangular, with a pair of median pits flanked proximally by a pair of lateral swellings. Pedipalpal trochanter not produced distally.

Tarsal-basitarsal spurs about 1/5 length of tarsus-basitarsus, claw missing. Tarsal-basitarsal segments of leg I of the following approximate proportions: 49-6-9-10-8-11-21. Other leg segment measurements given in Table 6.

Female unknown.

Variation.--Only one specimen of this species is known.

Remarks.--S^. drakos is placed in the simonis group on the

basis of the male's elongate abdomen and its possession of a

posterodorsal abdominal process, though not truncate as in most other males of this group, but round. That this species forms a distinct 106 complex with OTU #1 and OTU #2 is indicated by their geographic proximity and their morphological similarity. Further, these species may be closely related to species which Remy (1961) described from Surinam. The types of these species are not available for study, but his description leads me to believe that S^. surinamensis and S^. gladiator may belong in the simonis group.

Distribution.--This species is known only from Kartabo,

Bartica District, British Guiana.

Etymology.--Drakos is taken from the Greek word meaning dragon, a name which is inspired by the long abdomen of this species.

Schizomus simonis Hansen and Sorensen, 1905

(Figs. 91, 98, 113, 114; Tables 5, 6; Map 2)

Schizomus simonis Hansen and Sorensen, 1905:5, 7, 14, 15, 19, 22,

24, 38, 39, 42-44, 71-73; Chamberlin, 1922:12; Mello-Leitao,

1931:19; Giltay, 1935:7; Gertsch, 1940:3; Takashima, 1943:93;

Remy, 1961:504; Lawrence, 1969:219, 221, 223.

Cotypes.--An adult male, taken at St. Esteban, Venezuela, by

E. Simon (UZMK, examined); an adult female, taken at Colonia Tovar,

Venezuela, by E. Simon (UZMK, examined); an adult male and female

taken at Colonia Tovar, Venezuela, by E. Simon (NRS, examined).

Description.--Male. Color greenish. Carapace with two pairs

of dorsal and two apical setae. Eyespots oblong, indistinct.

Anterior sternum with 11 bifid setae. Abdominal terga I-VII with

two setae, terga VIII-IX with four setae, abdominal segments X-XII 107 elongate, tapering, segment XII with slight development of posterodorsal process. Vestigial stigmata nearly indistinguishable from sterna. Flagellum extended distally, apex acute; dorsal surface with two lateral swellings distally undercut by lateral pits.

Pedipalpal trochanter produced very slightly. Tarsal-basitarsal spurs about 1/6, claw about 1/4 length of tarsus-basitarsus.

Tarsal-basitarsal segments of leg I of the following approximate proportions: 33-5-6-7-9-8-14. Other leg segment measurements given in Table 6.

Female. Abdomen not elongate. Flagellum with four articles.

Six to eight pairs of spermathecae of varying size.

Comparisons,--See comparisons under S^, trinidanus and S^, acrocaudatus.

Distribution,--This species is known from Colonia Tovar and

St, Esteban, Venezuela,

Remarks,--The female's spermathecal arrangement is unique within the group. It is possible, however, that the female of this species has been misassociated with the male and that this is the female of another species. It may be noted in other species that the measurements of the legs are generally the same or fairly proportional between the sexes. This is not at all the case in S^. simonis, which

leads me further to believe the sexes may be mismatched in this species.

Variation.--The figure of the male's flagellum by Hansen and

Sorensen (1905) is much longer than that of the cotype available to me. 108

Schizomus trinidanus, new spo( ies

(Figs. 94, 99, 101, 116-118; Tables '., 6; Map 2)

Holotype.--An adult male, taken in Anma Valley, 8-1200 ft.,

Trinidad, during 10-22 February 1964, by P. Wvjodzinski (MCZ),

Allotype.—An adult female, taken with the holotype (MCZ).

Paratypes,--An adult male and eight >uiult females, taken with the holotype (MCZ),

Description.—Male, Color brownish. Carapace with two pairs

of dorsal and two apical setae, Eyespots in-<>gular, distinct.

Anterior sternum with 11 bifid setae. Abdominal terga I-VII with two

setae, terga VIII-IX with four setae, abdominal segments X-XII

slightly elongate, segment XII with truncate iiosterodorsal process.

Vestigial stigmata darker than sterna. Flagellum lanceolate, with a

pair of median depressions flanked proximal1\ by pair of lateral

swellings, Pedipalpal trochanter not produce-.-, apically, tarsal-

basitarsal spurs about 1/4, claw about 1/2 K-ngth of tarsus-basitarsus.

Tarsal-basitarsal segments of leg I of the following approximate

proportions: 30-5-6-6-7-7-11, Other leg segi-t>nt measurements given

in Table 6,

Female, Abdomen not elongate. Flagc'lum with four articles.

Lateral and median spermathecae short, nearly oqual sized, terminating

in slight sclerotized bulbs.

Comparisons,--Males of S^, trinidanus . like S^, simonis, have

only the pygydial segments of the abdomen elo-.^ate, but the eyespots

are more distinct in S, trinidanus, the post.-'odorsal abdominal 109 process is truncate in S^. trinidanus and is round in S^, simonis. If the female of S^. simonis is correctly assigned, then the longer flagellum in the latter is diagnostic. The multiple spermathecae of

S^. simonis also contrasts greatly with the two pairs found in other members of the group. The single dorsal depression of the male's flagellum in S^, tobago and the long spermathecae of the female will distinguish it from S^, trinidanus. The male of S^, acrocaudatus, which was collected with types of S^, trinidanus, can be distinguished by the very long flagellum and abdomen, the latter involving abdominal segments VII-XII, Unknown variability in the latter characters may leave the deep dorsal depressions more consistantly useful in distinguishing these two species.

Distribution,--This species is known from Arima Valley, Simla, and St, Augustine, Trinidad,

Etymology.--The specific name is an adjectival form taken from the type locality.

Variation,--The spermathecal morphology seems to be fairly consitant even from several localities. The specimen from Arima

Valley shows slightly less distinct apical bulbs, apparently because they are somewhat thicker basally. The three males from St. Augustine show only very slight elongation of the pygydial abdominal segments. This variation in secondary sexual characteristics is common in other species.

Additional records.--Simla: five females, bamboo debris, 26

April 1964, Chickering (MCZ); one female, 23 April 1964, Chickering 110

(MCZ); two females, 18 April 1964, Chickering (MCZ); one female,

20-21 October 1964, Chickering (MCZ); two females, 25 April 1964,

Chickering (MCZ); five females, 16 April 1964, Chickering (MCZ); four females, 28 April 1964, Chickering (MCZ); three females, 12

April 1964, Chickering (MCZ); three females, 19 April 1964,

Chickering (MCZ). Arima Valley: one female, 8-1200 ft., 10-22

February 1964, J. Rosen, P. Wygodzinsky (AMNH); four females, two immatures, 8-1200 ft,, 10-22 February 1964, J. Rosen, P, Wygodzinsky

(AMNH); one female, 10-22 February 1964, J, Rosen, P. Wygodzinsky

(AMNH), St, Augustine: three males, ten females, Weber (MCZ),

Schizomus acrocaudatus, new species

(Figs, 88, 97; Tables 5, 6; Map 2)

Holotype,--An adult male, taken at St, Augustine, Trinidad, by N. A. Weber (MCZ).

Description.--Male. Color brownish. Carapace with two pairs of dorsal and two apical setae, length 1.14. Eyespots irregular, indistinct. Anterior sternum with 10 setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae, abdominal segments

VII-XII slightly elongate, segment XII with small, truncate posterodorsal process. Vestigial stigmata darker than sterna.

Flagellum extremely long and distally very thin, a pair of median deep pits flanked laterally by pair of swellings, length .86, width .29. Pedipalpal trochanter not produced distally. Tarsal- basitarsal spurs about 1/5, claw about 1/3 length of tarsus-basitarsus. Ill

Tarsal-basitarsal segments of leg I missing. Female unknown.

Comparisons.--S^. acrocaudatus is very similar to S^. simonis males in respect to the development of the male's flagellum. The male cotype illustrated by Hansen and Sorensen (1905), though not the cotype available to me, shows the flagellum to be extremely long, attenuate, and indeed very similar to S^. acrocaudatus. The abdomen of S^. acrocaudatus is elongate from VII through XII, whereas the extension occurs in X through XII in ^. simonis. The flagellum of

S^. acrocaudatus is much thicker basally and the pair of dorsal depressions are clearly visible from directly above. The flagellum

of S^. simonis is flatter dorsally and the pair of depressions are not

clearly visible from above.

Distribution,--This species is known only from a single male

from St. Augustine, Trinidad.

Etymology.--The specific name is taken from the Latin acr-

meaning sharp and caud- meaning tail, describing the morphology of

the male's flagellum.

Remarks.--As in other species with elongate abdomen and

flagella, the degree of elongation can he highly variable. It is not

safe to consider the extreme elongation of the flagellum of this

species as being characteristic, nor diagnostic, rather the basal

configuration of elevations and depressions. 112

Schizomus flavescens Hansen and Sorensen, 1905

(Fig. 112; Tables 5, 6; Map 2)

Schizomus flavescens Hansen and Sorensen, 1905:39, 44-46, 47, 73;

Mello-Leitao, 1931:17; Hilton, 1933:92; Giltay, 1935:6;

Takashima, 1943:93.

Cotypes.--An adult female, taken at Corosul, near Caracas,

Venezuela, by E, Simon (UZMK, examined); two adult females and an immature, taken with the female above (MNHN); an adult female, taken at Corosul, near Caracas, Venezuela, in 1888, by E, Simon (NRS, examined).

Description.--Male unknown. Female as follows. Color brownish. Carapace with two pairs of dorsal and two apical setae.

Eyespots indistinct. Anterior sternum with 11 bifid setae.

Abdominal terga I-VII with two setae, terga VIII-IX with four setae.

Vestigial stigmata not distinguishable from sterna, Flagellum with four articles, extremely long. Pedipalpal trochanter produced slightly distally. Tarsal-basitarsal spurs about 1/4, claw about

1/2 length of tarsus-basitarsus. Tarsal-basitarsal segments of leg I of the following approximate proportions: 43-8-7-8-9-11-22. Other leg segment measurements given in Table 6,

Comparisons,—The female of this species is easily distinguished from others of the simonis group by the extremely long flagellum. The species most closely approaching S_, flavescens in length of female's flagellum is S^, simonis in which the flagellum of the cotype is ,52 113 as opposed to .61 in S^. flavescens. The morphology of the spermathecae is most similar to and, considering unknown possible variation, perhaps not safely distinguishable from S^, tobago and

S^. centralis. The extremely long first legs also distinguish it from S^. tobago, but tend to ally S^. flavescens with S^. drakos which is known, hov;ever, only by a male. The dorsal carapacal setation of the female of S^. drakos, however, probably consists of three pairs of dorsal setae as in the male.

Distribution,--This species is known only from Corosul,

Venezuela,

Remarks,--Hansen and Sorensen (1905) report a close similarity of this species to females of S^, dispar. The latter species was, however, not available for my studies,

Schizomus tobago, new species

(Figs, 87, 100; Tables 5, 6; Map 2)

Holotype.--An adult male taken in Tobago, during April 1916, by Th(omas) M(ortensen) (UZMK),

Allotype.—An adult female taken with the holotype (UZMK).

Description,--Male, Color brownish. Carapace with two pairs of dorsal and two apical setae. Eyespots indistinct. Anterior sternum with 11 bifid setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segments VII-XII elongate, segment XII with slight truncate posterodorsal process. Vestigial stigmata not distinguishable from sterna. Flagellum elongate, lanceolate, with a 114 single median depression undercutting a pair of lateral ridges.

Pedipalpal trochanter produced slightly apically, Tarsal-basitarsal spurs about 1/5, claw about 1/2 length of tarsus-basitarsus,

Tarsal-basitarsal segments of leg I of the following approximate proportions: 36-6-7-7-8-8-18, Other leg segment measurements given in Table 6.

Female. Abdomen not elongate. Flagellum composed of four articles. Spermathecae nearly equal sized, long, terminating in sclerotized bulbs.

Comparisons.--S^. tobago is similar to several species in the simonis group. The male's flagellum is most similar in basal form to S^. mumai and S^. centralis in having a single median depression.

It is, in fact, very similar in form to S^. mumai, but S^. centralis males have much shorter flagella. The flagella of S^. simonis and

S. trinidanus are also similar in form to S^. tobago, but the former

species have a pair of median dorsal depressions. The female's

spermathecae in S^, tobago, S^, flavescens, and S^, centralis are all

very similar, and perhaps not safely distinguishable, S^, mumai females have lateral spermathecae twice as long as the medians,

whereas in S^. tobago they are about equal in size.

Distribution,--This species is known only by a single

collection from Tobago,

Etymology,--The specific name is a noun used in apposition. 115

Schizomus mumai, new species

(Figs. 92, 93, 96, 107-109; Tables 5, 6; Map 2)

Holotype.--An adult male, taken at Coto, Costa Rica, on 28

June 1957, by E. Dixon (AMNH),

Allotype,--An adult female taken at Coto, on 11 September

1957, by E. Dixon (AMNH),

Paratypes.--An adult male taken with the holotype (AMNH); two adult females taken at Coto, on 19 July 1957, by E. Dixon (AMNH); two adult females taken at Coto, on 13 June 1957, by E. Dixon (AMNH); an adult female, taken at Golfito, on 17 September 1957, by E. Dixon

(AMNH); an adult female, taken at Coto, on 4 September 1957, by E.

Dixon (AMNH).

Description.--Male. Color brownish. Carapace with two pairs of dorsal and two apical setae. Eyespots, irregular, but distinct.

Anterior sternum with 11 bifid setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segment XII with well developed, truncate posterodorsal process, segments V-XII elongate. Vestigial stigmata slightly darker than sterna, Flagellum elongate, with a median depression flanked by a pair of proximal elevations, Pedipalpal trochanter slightly produced distally, tarsal-basitarsal spurs about

1/7, claw about 1/2 length of tarsus-basitarsus, Tarsal-basitarsal segments of leg I of the following approximate proportions: 45-7-7-7-

7-9-20, Other leg measurements given in Table 6, 116

Female, Abdomen not elongate, Flagellum composed of four articles. Lateral spermathecae about twice as long as medians, both pairs terminating in sclerotized bulbs, the laterals being much the larger,

Comparisons.--S^, mumai is most similar to S^, centralis in both sexes. In the few males available for comparison, the abdomen is consistantly less attenuated in S^, centralis, involving segments

VII-XII, while in S^. mumai the attenuation involves segments V-XII.

This is probably, however, a variable characteristic in both species.

The flagella of the males are distinct, though variable within the species, S^. mumai males have elongate flagella, whereas flagella of

S^. centralis males are relatively shorter. Flagellar lateral swellings are well defined in S^. mumai, but are undeveloped in S^. centralis. The spermathecae of the species are also similar; however, the laterals are twice as long as the medians in S_. mumai and only slightly longer than the medians in S_. centralis. The males of S_. mumai are most easily distinguished from males of other species known from within its range by its elongate abdomen. Females of S^. mumai are similar to S^. cubanicoides females, of the cubanicus group, but can quickly be distinguished by the number of pairs of dorsal carapacal setae; there are four pairs in S^. cubanicoides and two pairs in S^. mumai.

Distribution.--This species is known from Coto and Golfito,

Costa Rica. 117

Etymology.--The specific name is a patronym given for Martin

H. Muma, who first recognized this species as distinct from existing taxa.

Variation.--The flagellum of the males is considerably different in the two specimens available, the holotype's being very much longer than the paratype's. While the length is different, there seems to be only a very slight difference in the basal portion, or body, of the flagellum.

The female's spermathecae show considerable variation even at the same locality. The sclerotized bulbs all seem to be approximately similar, though the spermathecal tubes vary in configuration.

Schizomus centralis Gertsch, 1941

(Figs, 90, 95, 110, 111; Tables 5, 6; Map 2)

Schizomus centralis Gertsch, 1941:13-14,

Holotype,--An adult male, taken on Barro Colorado Island,

Panama Canal Zone, on 19 July 1938, by E, G, Williams (AMNH, examined)

Allotype,--An adult female, taken on Barro Colorado Island,

on 20 July 1938, by E, G, Williams (AMNH, examined).

Description,--Male, Color brownish. Carapace with two pairs

of dorsal and two apical setae, Eyespots oval, distinct. Anterior

sternum with 11 bifid setae. Abdominal terga I-VII with two setae,

terga VIII-IX with four setae, segments VII-XII elongate, tapering,

segment XII with well developed, truncate posterodorsal process. Figs, 87-94,--Dorsal views of male's flagella of the simonis group: 87, ^. tobago; 88, S^. acrocaudatus; 89, S^, drakos; 90, S^. centralis; 91, S^. simonis; 92, 93, S^. mummai; 94, £. trinidanus.

Scale: Figs, 87, 89, 92, 93, 1 cm = ,08 mm; Figs, 88, 90, 91, 94,

1 cm = ,06 mm.

Figs, 95-106.--Parts of schizomids of the simonis group: 95-100,

lateral views of male's flagella: 95, S^, central is; 96, S^. mummai;

97, S^, acrocaudatus; 98, S^, simonis; 99, S^, trinidanus; 100, S^. tobago;

101, lateral view of female's flagellum of S^, trinidanus; 102, 103, dorsal views of male's posterodorsal abdominal process: 102, S^,

centralis; 103, S^, mummai; 104-106, female's spermathecae: 104, 105,

OTU #1. Scale: Figs. 95, 99, 101-103, 1 cm = ,06 mm; Figs. 96-98,

100, 1 cm = ,12 mm; Figs, 104-106, 1 cm = ,04 mm.

Figs. 107-118,—Female's spermathecae of the simonis group:

107-109, S_, mummai: 107, from Golfito; 108, 109, from the type locality; 110, 111, S^, centralis; 112, S^, flavescens; 113, 114, S. simonis: 113, view from above the perpendicular; 114, viewed from the perpendicular; 115, S^. tobago; 116-118, S^. trinidanus from various localities: 116, St. Augustine; 117, Simla; 118, the type locality. Scale: 1 cm = ,04 mm. 109

114 Table 6,—Measurements of the members of the simonis group:

1, one female, OTU #1; 2, one female, OTU #2; 3, one male, S^, drakos;

4, one male, S^. simonis; 5, one female, S^. simonis; 6, three males,

S^. trinidanus; 7, three females, S^, trinidanus; 8, one female, S^, flavescens; 9, one male, S^, tobago; 10, one female, S^, tobago; 11, two males, S^, mummai; 12, three females, S_, mummai; 13, three males,

5, centralis; 14, five females, S, centralis. 125

TABLE 6

Taxa n 1 2 3 4 5 6 7 8 1 1,08 1,29 1.34 .95 1.40 1,07-1,07 1.11-1.14 1.33

2 - ,38 .58 .40 .52 ,43-,45 ,37-,40 .61

3 - - ,38 ,19 - ,22-,23 - - 4 1.11 1,17 1,78 1,15 1.40 1,03-1.10 1,02-1,05 1.48 5 1.30 1,41 2,24 1.44 1.68 1.28-1,35 1,25-1.28 1.78 6 .96 1,06 1.64 1,06 1.22 ,93-,97 .93-,93 1.43 7 .80 .89 1,14 ,86 1.00 ,79-,82 .78-.78 1,08

8 .75 .82 1,04 ,65 .96 ,67-.71 .70-,72 1,00 9 .42 .49 .53 .33 .50 ,39-,40 .41-.44 ,61 10 ,45 ,50 ,65 ,40 ,55 ,39-,43 ,42-,43 ,65 n ,43 ,44 .56 .40 .56 .36-,38 ,37-,37 54 12 ,56 ,70 ,86 ,56 .42 ,58-,60 ,61-.62 .87 13 .31 .35 .37 ,23 ,40 ,25-.26 .28-,29 .42 14 .35 .39 ,49 .26 ,41 .30-.32 .30-,32 .45

15 .45 .47 ,63 .40 - ,35-,36 ,37-,40 .56

16 1.06 1.14 1.41 1.00 1.32 .95-.99 ,97-1,06 .89 17 ,46 ,55 .58 .36 .65 .44-.47 .47-.52 .40 18 ,71 .79 ,93 ,65 .89 .61-.63 ,64-,67 .49 19 ,64 .70 .90 ,53 .83 .54-.57 ,55-.58 .54 126

TABLE 6 (continued)

Taxa n 9 10 11 12 13 14 1 1,23 1,12 1.07-1.24 1,07-1,14 ,92-1,10 1.03-1.12 2 .79 ,41 .59-.87 ,39-,41 ,44-,45 .35-.37

3 ,22 - .23-.23 - ,25-,25 - 4 1,32 1,00 1.05-1.34 ,91-,94 1,00-1,28 .89-.98 5 1,67 1,23 1.31-1.72 1,10-1,15 1,22-1,65 1.05-1.15 6 1.25 ,89 .96-1.23 ,83-.85 .91-1.15 .81-.84 7 .90 ,72 .84-,95 .76-.77 .82-,93 .70-.76 8 .77 .65 .68-.83 .65-.67 .63-,80 ,58-,69

9 .40 .41 ,41-.51 .40-.41 .37-,47 .38-.42 10 ,52 ,40 ,44-,58 .42-.44 .39-.53 .38-.40

11 ,43 ,35 .37-,46 .35-,38 ,35-.36 ,34-,38

12 .68 ,61 ,59-,72 .59-,61 ,55-,65 ,58-,60

13 .31 .27 ,26-,35 ,25-.28 ,25-,35 .26-.28

14 .37 ,30 ,30-,38 ,32-,33 ,28-,36 .28-.31

15 .43 .36 ,37-,44 ,34-.37 ,34-,40 .35-.38 16 1,09 ,93 .93-1.03 .89-.93 .93-1.06 ,91-.96 17 ,55 .45 .49-.58 .43-.48 ,46-.52 ,45-,49

18 ,69 .61 ,64-.76 ,63-,65 .60-.75 ,59-,65

19 .58 .54 ,55-,69 ,54-,55 .52-.65 .53-,56 127

Vestigial stigmata slightly darker than sterna, Flagellum lanceolate, with a distal median depression undercutting a more proximal ridge,

Pedipalpal trochanter produced only slightly, tarsal-basitarsal spurs about 1/6, claw about 1/2 length of tarsus-basitarsus,

Tarsal-basitarsal segments of leg I of unknown proportions.

Female, Abdomen not attenuate, Flagellum composed of four articles. Lateral spermathecae somewhat longer than medians, both terminating in sclerotized bulb of nearly equal size.

Comparisons.--This species is most closely related to S^. mumai. For comparisons see the treatment under that species.

Distribution.--This species is known only from Barro

Colorado Island, Panama Canal Zone.

Remarks.--The holotype lacks the flagellum, abdominal segment XII and nearly all its appendages.

Additional records,--Barro Colorado Island, Panama Canal

Zone: one male, 20-24 June 1924, N, Banks (MCZ); four females,

20-21 May 1964, Chickering (MCZ); two females, July 1969, S.

Lawrence, B, and T. Hlavac (MCZ); one male, 1943-1944, J. Zetek

(MCZ); one female, K, W. Cooper (AMNH).

The brasiliensis group

Members of this group may be small to large sized (.91 - 1.48 carapacal length). The color is brownish to greenish. The eyespots are always distinct, but may vary in shape from irregular to oval to round. The carapace has three or four pairs of dorsal and two apical 128 setae. The abdomen is never attenuated. Males have a very slightly to well developed posterodorsal abdominal process, which can be round, bifid, or truncate apically. The male's flagellum is usually large and nearly globose, with often elaborate dorsal modifications.

The female's flagellum is usually moderate in length (.28), but may be long in large species (.48), and is composed of three articles.

The females usually have two pairs of spermathecae, usually of similar size. The apical portions are sometimes extremely large, almost circular, and highly sclerotized, whereas other individuals have considerably smaller unsclerotized bulbs. The pedipalps are usually highly dimorphic, but variably so. A slight elongation is usually manifest, but more often the segments are heavily developed.

The trochanter is noticeably produced; the femur sometimes has a mesal tooth, as does the patella.

Distribution.--Mexico: Oaxaca, Tabasco, Chiapas. Central

America: Costa Rica. South America: Columbia, Ecuador, Brasil,

Bolivia.

Subordinate taxa.--Trilobatus complex: S^. stewarti, S^.

trilobatus, S^. lacandonus; brasiliensis complex: S^. cuenca, S^,

sturmi, S_, brasiliensis, OTU #7, OTU #8, OTU #9, OTU #10, OTU #11;

gaXlipatellatus lineage within the brasiliensis complex: OTU #12,

S.- macarensis, S^, cumbalensis, S^, pal lipatellatus. Table 7.--Comparisons of the members of the brasiliensis group. For explanation of the characters see Phylogeny. 130

4-) M fll 3 0.-P 1- (O

Id in cns- OJ (0

OJ ra J- r-

cnt-

0)

S- 3 o

0) to a (u

I V) C 3 10 C O O 10 T3

S^ (d i. I—

< LU UJ o 5:1- CO CD < O Uj I— a LXJ HH 2i S3 t:3 i: I- Si Maps 6-7,—Distribution of schizomids of the briggsi and

brasiliensis groups: Map 6, briggsi group members: 1, S^. briggsi;

2, S^. shoshonensis; 3, S^. belkini; 4, S^. pentapeltis; 5, S^. joshuensis; 6, ^. borregoensis; 7, S^. wessoni. Map 7, brasiliensis group members: 1, S^. stewarti; 2, S^. trilobatus; 3, S^. lacandonus;

4, S.. pal lipatellatus; 5, OTU #9; 6, S^. sturmi; 7, S^. macarensis;

8, OTU #8; 9, ^. brasiliensis; 10, S.. cumbalensis; 11, OTU #12; 12,

OTU #10; 13, S. cuenca; 14, OTU #11; 15. OTU #7. 132 133

Schizomus stewarti Rowland, 1973

(Figs. 119, 130; Tables 7, 8; Map 7)

Schizomus stewarti Rowland, 1973:139-140.

Holotype.—An adult male, taken in Cueva del Guayabo, 12 km

NE Valle Nacional, Oaxaca, Mexico, on 29 December 1972, by J.

Reddell, D. McKenzie, M. McKenzie, and S. Murphey (AMNH, examined).

Paratype.--An immature, taken with the holotype (AMNH, examined).

Description.--Male. Color greenish. Carapace with three pairs of dorsal and two apical setae. Eyespots distinct, irregular.

Metapeltidium entire. Anterior sternum with nine bifid setae.

Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segment XII with a slight development of posterodorsal process.

Vestigial stigmata darker than sterna. Flagellum vaguely trilobate, with a pair of slight median elevations. Pedipalpal trochanter produced acutely apically, tarsal-basitarsal spurs about 1/5, claw about 1/2 length of tarsus-basitarsus. Tarsal-basitarsal segments of leg I of unknown proportions. Other leg segment measurements given in Table 8.

Female unknown.

Comparisons.--This species stands quite by itself within this group in morphology of the male's pedipalp and flagellum. It most closely resembles males of S^. moisii in development of the flagellum, but the latter completely lacks the posterodorsal abdominal process. 134

Distribution.--This species is known only from Cueva del

Guayabo, near Valle Nacional, Oaxaca, Mexico.

Remarks.--As the most primitive member of the brasiliensis group this species shares plesiomorphic characters with primitive members of other groups. It is important to use the complete diagnostic combination of characters in order to properly place this and other species in their proper groups since single, key synapomorphic character states are often lacking in primitive taxa.

Schizomus trilobatus Rowland, 1975

(Figs. 120, 131, 146; Tables 7, 8; Map 7)

Schizomus trilobatus Rowland, 1975:11-13.

Holotype.--An adult male, taken in Las Grutas del Cocona,

Tabasco, Mexico, on 24 July 1973, by J. M. Rowland and J. R. Reddell

(TTU, examined).

Allotype.--An adult female, taken with the holotype (TTU, examined).

Paratypes.--An adult male and five adult females taken with the holotype (AMNH, examined).

Description.--Male. Color greenish. Carapace with three pairs of dorsal and two apical setae. Eyespots distinct, irregular.

Metapeltidium entire. Anterior sternum with 10 bifid setae.

Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segment XII with slightly rounded development of posterodorsal process. Vestigial stigmata darker than sterna, Flagellum strongly 135 trilobate, with a pair of medial depressions, Pedipalpal trochanter produced distally, tarsal-basitarsal spurs about 1/5, claw about 1/2 length of tarsus-basitarsus. Tarsal-basitarsal segments of leg I of the following approximate proportions: 42-8-6-8-7-8-15, Other leg measurements given in Table 8,

Female. Flagellum composed of three articles. Spermathecae composed of a single pair, highly sclerotized, large, on narrow stalks.

Comparisons.--See under S^. lacandonus.

Distribution.--This species is known only from Grutas del

Cocona, Tabasco, Mexico.

Remarks.--Grutas del Cocona contains at least two species of schizomids. S^. trilobatus has been collected from the twilight zone and apparently does not extend into remote areas of the cave occupied by S^, pecki, of the pecki group. The latter species is troglobitic and apparently is excluded from the twilight zone by S^. trilobatus, an obvious invader from outside the cave environment.

Schizomus lacandonus Rowland, 1975

(Figs. 121, 132, 139, 149; Tables 7, 8; Map 7)

Schizomus lacandonus Rowland, 1975:16-18.

Holotype,--An adult male, taken at Ruinas de Palenque, Chiapas,

Mexico, on 25 July 1973, by J, M, Rowland and J, R. Reddell (TTU,

examined). 136

Female,--An adult, taken at Ruinas de Palenque, Chiapas,

Mexico, on 6 July 1949, by C, J, Goodnight (AMNH, examined).

Description,--Male, Color greenish. Carapace with three pairs of dorsal and two apical setae. Eyespots distinct, oval.

Metapeltidium entire. Anterior sternum with nine bifid setae.

Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segment XII with gently rounded, slightly developed posterodorsal process. Vestigial stigmata darker than sterna, Flagellum triangular, with a pair of median depressions, Pedipalpal trochanter produced acutely apically, tarsal-basitarsal spurs about 1/5, claw about 1/2 length of tarsus-basitarsus, Tarsal-basitarsal segments of leg I of the following approximate proportions: 52-8-7-8-7-8-18,

Other leg measurements given in Table 8,

Female, Flagellum composed of three articles, Spermathecae composed of a single pair, highly sclerotized, large, on wide stalks.

Comparisons,--This species is closest to S^, trilobatus and shares with it several common characters. They are distinguished by size, S^. lacandonus being slightly larger (1,18 carapacal length) than

S_. trilobatus (1.09 carapacal length). The males are most easily distinguished by the flagellum, whcich is triangular in S^, lacandonus and distinctly trilobate in S^, trilobatus. Females of these species are very similar, but the spermathecal stalks are much wider in S^, lacajidonus^, and are more strongly sclerotized in S^, trilobatus.

Distribution.--This species is known only from Ruinas de

Palenque, Chiapas, Mexico. 137

Remarks.--S^. lacandonus is one of five species that occur

apparently sympatrically near Palenque, Chiapas, Other species

include an undescribed member of the pecki group, S^. infernalis, S^.

portoricensis, and another undescribed species recently brought to my attention by James R, Reddell, but not treated in this study.

These species are most easily distinguished on the basis of the

male anatomy, but can also be distinguished by external female

characters and the spermathecae. In the case of the last species

mentioned none of the anatomy has been studied closely, but it can

be noted here that the species can be easily distinguished by the

presence of multiple setae on the abdominal terga, a unique

development within the order.

Schizomus cuenca, new species

(Figs. 124, 134, 137, 142; Tables 7, 8; Map 7)

Holotype.--An adult male, taken in Cuenca, Ecuador, on 3

April 1942, by D. and H. Frizzell (AMNH).

Description.--Male. Color brownish. Carapace with three

pairs of dorsal and two apical setae. Metapeltidium split. Eyespots

oval, distinct. Anterior sternum with 10 bifid setae. Abdominal

terga I-VII with two setae, terga VIII-IX with four setae, segment

XII with well developed, truncate posterodorsal process. Vestigial

stigmata darker than sterna. Flagellum trilobate, with a deep median

depression, deeply sculptured. Pedipalpal trochanter produced acutely

apically, femur with a spur, patella curved, Tarsal-basitarsal spurs 138 about 1/6, claw about 1/2 length of tarsus-basitarsus, Tarsal- basitarsal segments of leg I of the following approximate proportions:

49-6-11-10-10-11-19, Other leg segment measurements given in Table 8,

Female unknown.

Comparisons.--S^, cuenca is most similar in the morphology of the male's flagellum to S^. sturmi, S^. pallipatellatus and S^, brasiliensis in that they are all trilobate, with large lateral lobes and deep dorsal impressions. S^. pallipatellatus has the most distinctly trilobate flagellum and a single median depression, while the other species have double depressions. The posterodorsal abdominal process of the species in question are rather distinctive.

In S^. pallipatellatus it is bifid, in S^. cuenca it is broadly truncate, and in S^. sturmi and S^, brasiliensis it is gently rounded.

The spermathecae of S^, pallipatel latus are similarly greatly expanded, while S^, sturmi females have them more greatly expanded. The female is not known in S^. cuenca and that of S^, brasiliensis was not studied.

Schizomus sturmi (Kraus), 1975 NEW COMBINATION

(Figs, 126, 136, 140, 143, 152; Tables 7, 8; Map 7)

Trithyreus sturmi Kraus, 1957:245, 247-249; Sturm, 1958:142-143;

Kraus and Beck, 1967:404-405; Rowland, 1972a:70,

Holotype,--An adult male, taken "Kolumbien: Hang am Ostrand der Hochebene von Bogota, etwa 3 km vom Stadtrand entfernt,

2800-3000 m" in November, 1955, April, September or October 1956, by H. Sturm (SMF #9818, examined). 139

Paratypes,--Six adult females and eight immatures taken with the holotype (SMF #9819-9821, not examined); one adult male taken with the holotype (AMNH, examined); two adult males, two adult females, and one immature taken with the holotype (H, Sturm's collection, not examined); one adult male, four adult females and one immature taken "Nahere Umgebung der Stadt, 2900 m" (SMF #9822, not examined); one adult female taken with the latter (AMNH, examined).

Description,--Male, Color brownish. Carapace with three pairs of dorsal and two apical setae, Metapeltidium split or entire.

Eyespots oval, distinct. Anterior sternum with 11 bifid setae.

Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segment XII with gently rounded, distinct posterodorsal process.

Vestigial stigmata darker than sterna, Flagellum semicircular, apically tilobate to truncate, with a pair of median depressions,

Pedipalpal trochanter produced apically, femur with a spur, tibia

curved, tarsal-basitarsal spurs about 1/5, claw about 1/2 length of

tarsus-basitarsus, Tarsal-basitarsal segments of leg I of the

following approximate proportions: 46-6-9-9-12-11-18. Other leg

segment measurements given in Table 8,

Female, Flagellum composed of three articles. Median and

lateral spermathecae similar, narrow basally, expanded to nearly

circular apically, with no concentration of sclerotization.

Comparisons.--See under S. cuenca. 140

Distribution,--This species is known from near Bogata,

Columbia,

Remarks,--Sturm (1958, 1973) has made considerable contribution to the knowledge of the behavior of schizomids in general in studies of this species,

Schizomus brasiliensis (Kraus), 1967 NEW COMBINATION

(Figs, 123, 129; Tables 7, 8; Map 7)

Trithyreus brasiliensis Kraus, in Kraus and Beck, 1967:401-404;

Beck, 1968a:248-249; Beck, 1968b:76-78; Rowland, 1972a:70;

Brignoli, 1973:3.

Holotype,—An adult male, taken "Brasilien (Amazonas): bei

Manaus, Reserva Ducke des I,N,P,A., Bachsenke oberhalb des

Accompamento, Urwald mit dichtem Unterholz aus Palmen" in January

or February, 1966, by L. Beck (SMF #11919, examined).

Paratypes.--Two adult females and two immatures, taken with

the holotype (SMF #12467, 12768, examined).

Description.--Male. Color brownish green. Carapace with

three pairs of dorsal and two apical setae. Eyespots irregular, but

distinct. Metapeltidium split. Anterior sternum with 11 entire

setae. Abdominal terga I-VII with two setae, terga VIII-IX with

four setae, segment XII with gently rounded, distinct posterodorsal

process. Vestigial stigmata slightly darker than sterna, Flagellum

trilobate, with a deep median pit, median and lateral swellings,

deeply sculptured, Pedipalpal trochanter extremely and acutely 141 produced, femur with a spur, patella curved, Tarsal-basitarsal spurs about 1/5, claw about 1/2 length of tarsus-basitarsus,

Tarsal-basitarsal segments of leg I of unknown proportions. Other leg segment measurements are given in Table 8,

Female. Flagellum composed of three articles, Pedipalps unarmed, Spermathecae not studied.

Comparisons.--See under S^. cuenca.

Distribution,--This species is known from near Manaus, Brasil,

Remarks,--Beck (1968a, 1968b) has made distributional and

behavioral reports on this species,

Schizomus sp,, OTU #7

(Fig. 154; Tables 7, 8; Map 7)

Specimens examined,--Three adult females and four immatures,

taken 7 km N Leticia, Amazonas District, Brasil, during 20-25

February 1972, by S, Peck (AMNH),

Description.--Male unknown. Female, Color brownish green.

Carapace with three pairs of dorsal and two apical setae,

Metapeltidium split. Anterior sternum with 11 bifid setae. Abdominal

terga I-VII with two setae, terga VII-IX with four setae. Vestigial

stigmata lighter than sterna, Flagellum composed of three articles,

Pedipalpal trochanter produced slightly, tarsal-basitarsal spurs about

1/5, claw about 1/2 length of tarsus-basitarsus, Tarsal-basitarsal

segments of leg I of the following approximate proportions:

27-4-4-5-6-5-14, Other leg segment measurements given in Table 8, 142

Median and lateral spermathecae about equal lengths, laterals somewhat expanded distally, medians expanded basally, no localized sclerotization.

Comparisons,--OTU #7 is distinct from the members of this group in that the base of the median spermathecae is much wider than the apex. OTU's #8-11 are very similar in most respects, but perhaps can be tentatively diagnosed on the basis of spermathecal characters used in combination with other characters, OTU #8 is most distinct among the remaining taxa in that it has very narrow spermathecae,

OTU #9 has slightly thickened apical spermathecal walls, as in OTU

#12, but the terminal bulbs are more distinct, OTU's #10 and #11 are very similar in development of the female's spermathecae, though the terminal bulb seems slightly more distinct in OTU #10, S^. cumbalensis is again very similar in development of the spermathecae to the latter species, though it is readily distinguishable by the white patella I,

OTU #10 is a much larger species than OTU #11, and has a disproportionately larger flagellum.

Remarks,--I have described several taxa in this group to which I am reluctant to give formal nominal status. These taxa are not represented by males and while some may be fairly distinct in morphology of the female's spermathecae, it has been shown that the intraspecific limits of variation in this character can be so great as to be only useful when used in conjunction with secondary sexual characters of the male, I feel that this study is served, however. 143 by a maximal utilization of these taxa in phylogenetic and zoogeographic interpretations, even though they are not given formal taxonomic status,

Schizomus sp,, OTU #8

(Fig. 145; Tables 7, 8; Map 7)

Specimen examined.--An adult female, taken at Santarem,

Taperinha, Brazil, on 29 October 1970, by S. L, Tuxen (UZMK),

Description,--Male unknown. Female, Color brownish green.

Carapace with three pairs of dorsal and two apical setae, f'tetapeltidium split, Eyespots oval, distinct. Anterior sternum with 11 bifid setae. Abdominal terga I-VII with two setae, terga

VIII-IX with four setae. Vestigial stigmata lighter than sterna,

Flagellum composed of three sections, Pedipalpal trochanter produced distally, tarsal-basitarsal spurs about 1/4, claw about

1/2 length of tarsus-basitarsus, Tarsal-basitarsal segments of leg

I of the following approximate proportions: 24-4-4-4-5-6-13, Other leg segment measurements given in Table 8, Both median and lateral spermathecae very similar in form and shape; slightly divergent, not expanded distally; unevenly sclerotized along most of the length.

Comparisons,--See under Schizomus sp,, OTU #7.

Distribution.--This taxon is known only from Santarem,

Taperinha, Brasil.

Remarks.--See under Schizomus sp., OTU #7. 144

Schizomus sp,, OTU #9

(Fig, 153; Tables 7, 8; Map 7)

Specimen examined,--An adult female, taken at El Saladito,

Valle, Columbia, on 29 August 1967, by P, Wygodzinsky (AMNH),

Description,--Male unknown. Female, Color brownish green.

Carapace with three pairs of dorsal and two apical setae,

Metapeltidium split, Eyespots distinct, irregular. Anterior sternum with 11 bifid setae. Abdominal terga I-VII with two setae, terga

VIII-IX with four setae. Vestigial stigmata lighter than sterna,

Flagellum composed of three articles, Pedipalpal trochanter produced distally, tarsal-basitarsal spurs about 1/4, claw about 1/2 length of tarsus-basitarsus, Tarsal-basitarsal segments of leg I of the following approximate proportions: 34-4-6-6-7-7-14. Other leg segment measurements given in Table 7. Both median and lateral spermathecae of about equal size, medians terminate in bulb about twice diameter of those of laterals, both sclerotized, stalk of laterals about twice as wide as that of medians, expanded basally.

Comparisons,--See under Schizomus sp,, OTU #7,

Distribution,--This taxon is known only from El Saladito,

Valle, Columbia,

Remarks,--See under Schizomus sp., OTU #7, 145

Schizomus sp., OTU #10

(Fig, 147; Tables 7, 8; Map 7)

Specimens examined,--An adult female and an immature, taken at Banos, Ecuador, in April 1965, by J, and N. Leleup (MACB).

Description.--Male unknown. Female, Color brownish green.

Carapace with four pairs of dorsal and two apical setae,

Metapeltidium split, Eyespots irregular, distinct. Anterior sternum with 11 bifid setae. Abdominal terga I-VII with two setae, abdominal terga VIII-IX with four setae. Vestigial stigmata darker than sterna, Flagellum composed of three articles, Pedipalpal trochanter produced distally, tarsal-basitarsal spurs about 1/5, claw about 1/2 length of tarsus-basitarsus, Tarsal-basitarsal segments of leg I of the following approximate proportions:

39-4-7-7-8-8-15, Other leg segment measurements given in Table 8,

Median and lateral spermathecae small, similar, slightly smaller apically in the latter, but both expanded distally into small circular to oval terminal bulbs.

Comparisons,--See under Schizomus sp., OTU #7,

Distribution,--This taxon is known only from Banos, Ecuador,

Remarks,--See under Schizomus sp., OTU #7. 146

Schizomus sp., OTU #11

(Fig. 148; Tables 7, 8; Map 7)

Specimen examined.--An adult female, taken at Rio Benicito,

Chacoba, Bolivia, by W. J. Gertsch (AMNH),

Description.--Male unknown. Female, Color brownish green.

Carapace with three pairs of dorsal and two apical setae,

Metapeltidium split, Eyespots irregular, distinct. Anterior sternum with 11 entire setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae. Vestigial stigmata lighter than sterna. Flagellum composed of three articles. Pedipalpal trochanter produced slightly apically, tarsal-basitarsal spurs about 1/4, claw about 1/2 length of tarsus-basitarsus. Tarsal-basitarsal segments of leg I of the following approximate proportions: 26-4-5-5-5-6-14,

Other leg segment measurements given in Table 8, Median and lateral spermathecae very similar in shape and size, each gradually expanded distally, with no special sclerotization; medians convergent, laterals divergent.

Comparisons.--See under Schizomus sp., OTU #7,

Distribution,--This taxon is known only from Rio Benicito,

Chacoba, Bolivia.

Remarks.--See under Schizomus sp., OTU #7. 147

Schizomus sp., OTU #12

(Fig. 144; Tables 7, 8; Map 7)

Specimen examined.--An adult female, taken at Oriente Rio

Negro, Ecuador, in April 1965, by J. and N. Leleup (MACB).

Description.--Male unknown. Female. Color brownish green.

Carapace with four pairs of dorsal and two apical setae.

Metapeltidium split. Eyespots distinct, irregular. Anterior

sternum with 11 bifid setae. Abdominal terga I-VII with two setae,

terga VIII-IX with four setae. Vestigial stigmata lighter than

sterna. Flagellum composed of three articles. Pedipalpal trochanter

produced distally, tarsal-basitarsal spurs about 1/4, claw about 1/2

length of tarsus-basitarsus. Tarsal-basitarsal segments of leg I of

the following approximate proportions: 31-5-6-5-8-7-15, distal half

of patella white. Other leg segment measurements given in Table 8.

Median spermathecae about twice as long as laterals, medians slightly

convergent, terminating in a sclerotized bulb, laterals lightly

sclerotized along distal half, terminating in a slight bulf.

Comparisons,--See under S^, pallipatellatus.

Distribution,--This taxon is known only from Oriente Rio

Negro, Ecuador,

Remarks,--See under S^, pallipatellatus. 148

Schizomus macarensis (Kraus), 1957 NEW COMBINATION

(Figs, 127, 128; Tables 7, 8; Map 7)

Trithyreus macarensis Kraus, 1957:245, 249-250,

Holotype.—An adult male, taken "Kolumbien: Macarena,

Gebirgsstock am Fusse der Ostanden s, Villavicencio, nahe der

Mundung des Rio Zanza in den Rio Guejar, 400-500 m, in der Laubstreu eines primaren Hochwaldes" on 5 March 1956, by H, Sturm (SMF #9823, examined).

Paratypes,--An immature, taken with the holotype (SMF #9824, examined).

Description,--Male, Color brownish green. Carapace with three pairs of dorsal and two apical setae, Metapeltidium split,

Eyespots distinct, but irregular. Anterior sternum with 11 bifid setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segment XII with gently rounded, but distinct posterodorsal process. Vestigial stigmata darker than sterna.

Flagellum oval, with a pair of median depressions, united mesally, preceded by a distinct ridge. Pedipalpal trochanter distinctly and acutely produced distally, femur with a spur, tibia curved, tarsal-basitarsal spurs about 1/5, claw about 1/2 length of tarsus-basitarsus, Tarsal-basitarsal segments of leg I of the following approximate proportions: 33-5-5-5-7-7-18, distal half of patella white. Other leg segment measurements given in Table 8,

Female unknown. 149

Comparisons,--See under S^, pallipatellatus.

Distribution.--This species is known only from Macarena,

Columbia.

Remarks,--See under S^, pallipatellatus,

Schizomus cumbalensis (Kraus), 1957 NEW COMBINATION

(Figs, 125, 135, 151; Tables 7, 8; Map 7)

Trithyreus cumbalensis Kraus, 1957:245, 246-247; Kraus, 1967:402,

Holotype,--An adult male, taken "Sudkolumbien: Umgebung des

Ortes Cumbal, zwischen Pasto und Ipiales, 3100 m," on 30 June or 5

July 1956, by H, Sturm (SMF #9816, examined).

Paratypes,--Two adult females and four immatures (SMF #9817, examined), and one adult female and two immatures (collection of

Sturm, not examined), taken with the holotype.

Description,--Male. Color brownish. Carapace with three pairs of dorsal and two apical setae. Metapeltidium split. Eyespots distinct, oval. Anterior sternum with 11 bifid setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segment

XII with squarely blunt, well developed posterodorsal process.

Vestigial stigmata darker than sterna, Flagellum triangular, deeply sculptured apically, Pedipalpal trochanter produced extremely apically, femur with a spur, tibia curved, tarsal-basitarsal spurs about 1/6, claw about 1/2 length of tarsus-basitarsus, Tarsal-basitarsal segments of leg I of the following approximate proportions: 150

54-7-9-10-11-11-20, distal half of patella white. Other leg segment measurements given in Table 8,

Female, Flagellum composed of three articles. Median and

lateral spermathecae very small, weakly developed, slightly distally expanded and equal in size.

Comparisons,--See under S^, pallipatellatus,

Distribution,--This species is known only from Cumbal,

Columbia,

Remarks,--See under S^, pallipatellatus.

Schizomus pallipatellatus, new species

(Figs. 122, 133, 138, 141, 150; Tables 7, 8; Map 7)

Holotype.--An adult male, taken at Coto, Costa Rica, on

23 October 1957, by E, Dixon (AMNH),

Allotype,--An adult female, taken at Coto, on 12 September

1957, by E, Dixon (AMNH),

Paratypes,--An immature taken with the allotype (AMNH); an

adult female, taken at Coto, on 26 October 1957, by E, Dixon (AMNH);

an immature, taken at Coto, on 25 June 1957, by E, Dixon (AMNH); an

immature, taken at Coto, on 19 June 1957, by E, Dixon (AMNH); two

immatures, taken at Coto, on 19 July 1957, by E, Dixon (AMNH); an

adult female and an immature, taken at Golfito, on 17 June 1957, by

E. Dixon (AMNH),

Description,--Male, Color brownish green. Carapace with

three pairs of dorsal and two apical setae, Metapeltidium entire. 151

Eyespots distinct, round. Anterior sternum with 11 bifid setae.

Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segment XII with well developed, truncate posterodorsal process with a pair of lateral projections. Vestigial stigmata lighter than sterna, Flagellum trilobate, with a median depression flanked by a pair of whitish processes, Pedipalpal trochanter produced acutely apically, femur and patella with spurs, tarsal-basitarsal spurs about 1/5, claw about 1/2 length of tarsus-basitarsus, Tarsal- basitarsal segments of leg I of the following approximate proportions:

27-6-5-5-6-6-14, distal half of patella white. Other leg segment measurements given in Table 8.

Female. Flagellum composed of three articles. Median and lateral spermathecae distally expanded gradually, the laterals with subdistal constrictions, no areas of concentrated sclerotization.

Comparisons.--S^. pal lipatellatus is readily distinguished from other brasiliensis group members by possession of a white patella I, a distinctly trilobate male's flagellum, and large, distally expanded female's spermathecae. The flagellum is similar in form to that of S^. trilobatus, but S^, pallipatellatus has a single rather than double dorsal depression. The pedipalpal femur and patella of the latter, but not the former, have spurs. The males of OTU #12 is unknown, but males of the other species with patella I white are available, S^, macarensis, S_, cumbalensis, and i- pallipatellatus are quite diverse in shape of the male's flagellum.

The flagellum of S, cumbalensis is much more apically attenuate. r

Figs. 119-125,--Dorsal views of male's flagella of the brasiliensis group: 119, S^, stewarti; 120, S^, trilobatus; 121, S^, lacandonus; 122,

S^, pallipatellatus; 123, S^, brasiliensis; 124, S^, cuenca; 125, S^, cumbalensis. Scale: Figs, 119-123, 125, 1 cm = .06 mm; Fig. 124,

1 cm = .12 mm.

'^-^

Figs, 126-136,--Male's flagella of the brasiliensis group: 126,

127, dorsal views: 126, S^, sturmi; 127, S_. macarensis; 128-136, lateral views: 128, S^, macarensis; 129, S^, brasiliensis; 130, S_. stewarti; 131,

S^. trilobatus; 132, S^. lacandonus; 133, S^. pallipatellatus; 134, S. cuenca; 135, S^. cumbalensis; 136, S^. sturmi. Scale: Figs. 126, 135,

1 cm = .12 iwn; Figs, 127-133, 1 cm = ,06 mm; Figs, 134, 136, 1 cm-

,08 mm.

Figs, 137-145,--Parts of schizomids of the brasiliensis group:

137-140, lateral views of male's right pedipalps: 137, S^, cuenca; 138,

S^. pallipatellatus; 139, S_, lacandonus; 140, S^, sturmi; 141-143, dorsal views of male's posterodorsal abdominal process: 141, S^. pallipatellatus;

142, S^. cuenca; 143, S^. sturmi; 144-145, female's spermathecae: 144,

OTU #12; 145, OTU ^8. Scale: Figs. 137-143, 1 cm = .06 mm; Figs. 144,

145, 1 cm = .04 mm. 157 Figs. 146-154,—Female's spermathecae of the brasiliensis group:

146, S^, trilobatus; 147, OTU #10; 148, OTU #11; 149, i. lacandonus;

150, S^. pallipatellatus; 151, S^. cumbalensis; 152, S^. sturmi; 153,

OTU #9; 154, OTU #7, Scale: 1 cm = ,04 mm.

Table 8,--Measurements of the members of the brasiliensis group:

I, one male, S^. stewarti; 2, one male, S^, trilobatus; 3, one female,

S^, trilobatus; 4, one male, S^, lacandonus; 5, one female, S^, lacandonus;

6, one male, S^, cuenca; 7, one male, S^, sturmi; 8, one female, S^, sturmi; 9, one male, S^. brasiliensis; 10, one female, S^, brasiliensis;

II, one female, OTU #7; 12, one female, OTU #8; 13, one female, OTU #9;

14, one female, OTU #10; 15, one female, OTU #11; 16, one female, OTU

#12; 17, one male, S^. macarensis; 18, one male, S^. cumbalensis; 19. one female, S^. cumbalensis; 20, one male, S_. pallipatellatus; 21, one female,

^. pallipatellatus. 161

TABLE 8

Taxa

n 1 2 3 4 5 6 7 8 9

1 1.03 1.09 1.07 1,18 1.16 1,48 1,35 1.33 .95

2 .37 ,39 .24 .44 .28 ,57 .53 - ,38

3 .27 ,40 - .31 - ,54 .47 - ,36

4 - 1,50 1,03 1.73 .82 1.60 1,55 1.35 .93

5 - 2.02 1,26 2.31 1.12 1,93 1,90 1.50 1,15

6 - 1.52 .91 1.81 1.51 1,39 1,35 1,15 .80

7 - .91 .71 1.08 1,19 1,16 1,10 .98 ,73

8 .61 ,83 ,68 .90 .80 1.14 1.03 .93 ,62

9 .34 ,45 ,41 .50 ,43 .61 ,55 .45 .35

10 ,44 ,55 ,42 .64 ,48 ,68 .60 ,53 .35

11 ,41 ,54 .41 .53 .44 ,65 .57 ,55 ,35

12 ,51 .64 ,62 .76 ,68 ,95 .88 ,85 ,55

13 .25 ,26 ,30 .36 .31 ,48 .40 .35 ,25

14 .31 .37 ,31 .45 .33 ,50 .47 .43 ,30

15 ,40 ,48 ,40 .55 ,46 ,67 ,62 ,53 ,38

16 ,98 1,29 1,02 1.49 1,16 1,54 1.43 1.30 ,95

17 ,33 ,55 ,48 .62 ,49 ,76 .62 .55 ,46

18 ,71 .91 .70 1.07 ,80 1.00 .93 ,80 ,57

19 .55 ,79 ,61 .89 ,69 ,96 .85 ,78 ,50 162

TABLE 8 ( continued)

Tax a

n 10 11 12 13 14 15 16 17 18

1 1,04 .95 ,91 1.09 1.33 1.02 1,08 ,97 1.50

2 .25 .24 .20 .29 .38 .26 ,30 ,35 .66

3 ------.28 ,50

4 .85 ,76 .80 1.04 1,23 .91 1.07 1.05 1,55

5 1.00 ,88 ,93 1.12 1,45 1.03 1,26 1,28 1,83

6 .73 .62 .65 ,85 1,08 .72 ,92 ,95 1,38

7 .68 .65 ,60 .78 .88 .65 ,77 .80 1,20

8 .60 .52 .58 ,74 ,90 .68 ,74 .70 1,20

9 .25 .31 ,30 ,40 ,49 .38 ,42 .37 .62

10 .27 ,32 ,32 .41 ,53 .40 ,44 .40 .72

11 .32 .32 .33 .40 .50 .39 ,44 ,43 .70

12 .55 ,51 ,50 .62 .79 .59 ,65 ,60 1.05

13 .32 ,24 ,25 ,29 .41 .29 ,31 .36 .50

14 .33 ,24 ,25 ,33 .40 .31 ,32 ,30 ,57

15 ,35 ,33 ,33 ,40 .56 .41 ,45 ,43 .72

16 ,90 .87 .90 1,05 1.26 ,93 1.07 1,05 1.58

17 ,40 .36 .35 .48 .61 ,41 ,47 ,40 .70

18 .57 .52 .55 ,68 .84 ,63 ,71 .62 1,05

19 .50 ,51 .47 ,63 .81 ,58 ,66 ,57 .98 163

TABLE 8 (conti nued)

Taxa

n 19 20 21

1 1.48 .94 1,02

2 .47 .38 .26

3 - .49 -

4 1.35 .97 ,91

5 1.60 1,15 1,04

6 1.18 .82 .76

7 1.02 .69 .66

8 1.07 .63 ,66

9 .60 .28 ,33

10 .62 ,38 ,38

11 .60 ,35 .35

12 ,95 ,54 ,57

13 ,47 .25 ,25

14 .50 ,28 .30

15 ,62 ,37 ,35

16 1,45 ,91 ,93

17 ,68 .38 .41

18 .93 .60 .59

19 .87 ,53 .56 164 while S^. macarensis is nearly oval. The spermathecae of S^. cumbalensis are much smaller than those of S^. pallipatellatus, and much farther apart than in OTU #12, The spermathecae are more greatly expanded in S^, pallipatellatus than in OTU #12,

Remarks,--This and the preceding three species are tentatively treated together since they share the apomorphic character state of a white patella I, It is conceivable that a group with such a monothetic basis will divorce cladistically proximal species which have secondarily lost the key character. This assemblage of taxa, while certainly synapomorphic is not intended to represent a strictly monophyletic lineage.

The mexicanus group

Description,--Members of this group are moderate to large sized (.98-1,44 carapacal length). The color is usually brownish but green species occur. Eyespots are present in epigean species, but often are not evident in cave dwellers. They are usually indistinct, and ovoid with diffuse margins. The carapace has two or three pairs of dorsal and two apical setae. The male's abdomen

is never elongate. Males also lack the posterodorsal abdominal process. The male's flagellum is usually small, ovate and dorsally compressed, and the dorsal surface usually bears a pair of lateral depressions. The female's flagellum is short and composed of three articles. The female's spermathecae are generally characterized by

an enlargement of the median pair and reduction of the laterals. Table 9,—Comparisons of the members of the mexicanus group. For explanation of the characters see Phylogeny, 166

c0) ^ o ,_ T3 O +J ?<

c: -M

5 "

•s

•r- C •D'f- C 4J

-js: <: to E: a. i-i UJ E Li_ rD —1 X _i

P^. occidentalis; 2, P^, pachypalpus; 3, A, lucifer; 4, A^- huitzmolotitlensis, Map 4, circles represent pecki group members, triangles represent goodnightorum group members: 1, S_, firstmani;

2, S_, (cf,) sbordonii; 3, S^. lanceolatus; 4, OTU #8; 5, S^. pecki;

6, OTU #7; 7, OTU #2; 8, S_, orthoplax; 9, S_, guatemalensis; 10, S. silvino; 11, OTU #6; 12, S^. goodnightorum. Map 5, mexicanus group members: 1, S^. mulaiki; 2, S^. bartolo; 3, S^, davisi; 4, S_, mexicanus;

5, ^, lukensi; 6, S^, mitchelli; 7, S^, cookei; 8, S^, pallidus; 9, S_. moisii; 10, OTU #11; 11, OTU #1; 12, S, reddelli; 13, OTU #2; 14, unnumbered solid circles, and inset, S^. portoricensis. 168 169

Some species lack the laterals altogether, but in some they are nearly as large as the medians, A few species have terminal enlargements. The pedipalps sometimes are highly sexually dimorphic, but sometimes variably so. In species with dimorphic pedipalps an elaboration of all segments and production of a tibial spur apposible to the tarsus-basitarsus is characteristic; however, individuals with only slightly dimorphic pedipalps do not express the spur.

Distribution.--Excluding S^. portoricensis: United States:

Texas. Mexico: Nuevo Leon, Tamaulipas, San Luis Potosi, Guerrero,

Veracruz, Oaxaca, Chiapas, Campeche, Yucatan. Belize. Guatemala.

Si. portoricensis: Bermuda, Florida, southern Mexico, Central America, northern South America, Galapagos Islands.

Remarks,--S^, antillus Hilton, 1933, from Cuba has not been

studied by me. The types, reportedly deposited in the Pomona College,

California Museum, are apparently no longer available. It is my judgement from measurements given in the original description, that i. antillus is probably a junior synonym of S^, portoricensis, and is

so treated herein.

Subordinate taxa,--An unnamed complex: OTU #1, OTU #2; mexicanus complex: S^, mulaiki, S^. bartolo, S^. lukensi, S^. davisi,

i- reddel 1 i, S^. mexicanus; portoricensis complex: S_, pallidus, S^,

fiorton_censjs.' moisii complex: OTU #11, S_, moisii; mitchelli complex:

S. cookei, S. mitchelli. 170

Schizomus sp,, OTU #1

(Fig, 198; Tables 9, 11; Map 5)

Specimens examined,--An adult female and an immature, taken

in Grutas de Cacahuamilpa, Guerrero, Mexico, on 15 August 1966, by

J. Fish and J, Reddell (TTU, examined).

Description.--Male unknown. Female. Color brownish. Carapace with tv/o pairs of dorsal and two apical setae. Eyespots absent.

Anterior sternum with 11 bifid setae. Abdominal terga I-VIII with two setae, tergum IX with four setae. Vestigial stigmata darker than

sterna. Flagellum composed of three articles. Pedipalpal trochanter

produced distally, tarsal-basitarsal spurs about 1/4, claw about 1/2

length of tarsus-basitarsus. Tarsal-basitarsal segments of leg I of

the following approximate proportions: 43-5-8-10-11-10-20. Other leg

segment measurements given in Table 11. Median and lateral

spermathecae slightly divergent, the medians somewhat larger, medians

and laterals with variable, slightly sclerotized bulbs.

Comparisons,--See under Schizomus sp., OTU #2,

Distribution,--This taxon is known only from Grutas de

Cacahuamilpa, Guerrero, Mexico,

Remarks,--I have described three taxa in this group to which

I am reluctant to give formal nominal status. These taxa are not

represented by males and, while some may be fairly distinct in the

morphology of the female's spermathecae, it has been shown that the

intraspecific limits of variation in this character can be so great 171 as to be only useful when used in conjunction with secondary sexual characters of the male.

Schizomus sp,, OTU #2

(Fig. 194; Tables 9, 11; Map 5)

Specimens examined.--An adult female and an immature, taken

20 mi. S Juchatengo, 6000 ft., Oaxaca, Mexico, on 29 May 1971, by

S. Peck (TTU, examined).

Description.--Male unknown. Female, Color brownish.

Carapace with two pairs of dorsal and two apical setae. Eyespots distinctly triangular. Anterior sternum with nine entire setae.

Abdominal terga I-VII with two setae, terga VIII-IX with four setae.

Vestigial stigmata darker than sterna, Flagellum composed of three articles, Pedipalpal trochanter slightly produced distally, tarsal-basitarsal spurs about 1/4, claw about 1/2 length of tarsus-basitarsus, Tarsal-basitarsal segments of leg I of the following approximate proportions: 31-4-5-6-5-6-15, Other leg segment measurements given in Table 11, Median spermathecae about

1/4 longer than laterals, both pair straight, slightly divergent; each with apical portions sclerotized.

Comparisons.--OTU #2 is very similar to OTU #1, but is probably safely distinguished from the latter by its shorter flagellum.

Also, in OTU #1 the median spermathecae are outwardly divergent and have slightly thickened walls; in OTU #2 these are shorter, nearly straight and have greater thickening of the apical spermathecal walls. 172

Whether the latter spermathecal characters are reliable can only be known after more comparative work is done.

Distribution.--This taxon is known only from near Juchatengo,

Oaxaca, Mexico.

Remarks.--See under Schizomus sp., OTU #1.

Schizomus mulaiki Gertsch, 1940

(Figs. 162, 178; Tables 9, 11; Map 5)

Schizomus mulaiki Gertsch, 1940:1, 3-4; Rowland, 1971a:304.

Holotype.--An adult male, taken at Rio Grande City, Starr

County, Texas, on 21 June 1939, by S. Mulaik (AMNH, examined),

Paratype.--An adult male, taken at Edinburg, Hidalgo County,

Texas, on 2 June 1935, by S. Mulaik (AMNH, examined).

Description.--Male. Color brownish. Carapace with two pairs of dorsal and two apical setae. Eyespots indistinct. Anterior sternum with 12 bifid setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segment XII with no evidence of posterodorsal process. Vestigial stigmata darker than sterna,

Flagellum nearly circular, the body wider than long, with a distal median depression, Pedipalpal trochanter not produced distally, tarsal-basitarsal spurs about 1/6, claw about 1/3 length of tarsus-basitarsus. Tarsal-basitarsal segments of leg I of the following approximate proportions: 26-5-5-6-7-7-18. Other leg segment measurements given in Table 11.

Female unknown. 173

Comparisons,--S^, mulaiki is similar to other members of the mexicanus complex. It is distinct from the remainder of these, however, in having a small, yet distinct, single apical depression on the male's flagellum. Other males within the group have either no dorsal relief or have a pair of distal depressions.

Distribution.--This species is known from Hidalgo and Starr

Counties in southern Texas.

Remarks.--The morphology of the male's flagellum was probably arrived at by a central fusion of the ancestral pair of depressions.

A slight proximal cleavage of the pit is still apparent.

S^. mulaiki is known by only two collections of one specimen each; however, S. Mulaik (personal communication) stated that he has seen this species all along the Rio Grande, perhaps 100 miles northwest of the type locality.

Schizomus bartolo Rowland, 1973

(Figs. 158, 174, 193; Tables 9, 11; Map 5)

Schizomus bartolo Rowland, 1973a:6-7, 13-16, 18; Rowland, 1973c:135,

137.

Holotype.--An adult male, taken in Grutas de San Bartolo, 10 mi. SW Monterey, Nuevo Leon, Mexico, on 21 June 1969, by S. and J.

Peck (AMNH, examined).

Allotype,--An adult female, taken with the holotype (AMNH, examined). 174

Paratypes,--Eight immatures taken with the holotype (AMNH, examined); four adult females, taken in Grutas de San Bartolo, in

September 1971, by T, Raines (TTU, examined).

Description,--Male, Color pale brown. Carapace with three pairs of dorsal, the medians being very small, and two apical setae. Eyespots absent. Anterior sternum with 11 bifid setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segment

XII with no evidence of posterodorsal process. Vestigial stigmata darker than sterna. Flagellum globose, with no dorsal relief.

Pedipalpal trochanter produced distally, tarsal-basitarsal spurs about 1/4, claw about 1/2 length of tarsus-basitarsus. Tarsal- basitarsal segments of leg I of the following approximate proportions:

38-7-7-7-10-9-20. Other leg segment measurements given in Table 11.

Female. Flagellum composed of three articles. Median spermathecae large, straight, wide, slightly convergent; laterals smaller, convergent and constricted into divergent curves apically, the median apically sclerotized.

Comparisons.--See under S^. lukensi,

Distribution,--This species is known only from Grutas de San

Bartolo, Nuevo Leon, Mexico.

Additional records.--Nuevo Leon: Grutas de San Bartolo, four females and four immatures, February 1966, B. Russell (TTU), 175

Schizomus lukensi Rowland, 1973

(Figs, 160, 173, 191, 192; Tables 9, 11; Map 5)

Schizomus lukensi Rowland, 1973a:136-137, 138,

Holotype.--An adult male, taken in Cueva del Agua, 30 mi. SW

Soto la Marina, Tamaulipas, Mexico, on 31 October 1970, by W.

Russell, G, and J, Ediger (AMNH, examined).

Allotype,--An adult female, taken with the holotype (AMNH, examined).

Paratypes,--Two adult males and females, taken with the holotype (AMNH, examined).

Description.--Male. Color pale brownish. Carapace with three pairs of dorsal and two apical setae, Eyespots absent. Anterior sternum with nine bifid setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segment XII with no evidence of posterodorsal process. Vestigial stigmata lighter than sterna.

Flagellum lanceolate, with no dorsal relief. Pedipalpal trochanter produced distally, tarsal-basitarsal spurs about 1/5, claw about 1/2 length of tarsus-basitarsus. Tarsal-basitarsal segments of leg I of the following approximate proportions: 41-7-9-8-9-10-22. Other leg segment measurements given in Table 11.

Female. Flagellum composed of three articles. Spermathecae with medians extremely long, divergent, with slight terminal bulbs, only one very small lateral present. 176

Comparisons.--S^. lukensi is generally similar to other members

of the mexicanus complex. It shares with S^. bartolo the possession of

three pairs of dorsal carapacal setae and lack of dorsal relief on the male's flagellum. The male's flagellum is longer and narrower,

however, in S^. lukensi, The female's spermathecae are distinguishable

in that S^, lukensi is either missing or has very small lateral

spermathecae, whereas these are very well developed in S^. bartolo.

Distribution.--This species is known from Cueva de los

Cuarteles and Cueva del Agua, Tamaulipas, Mexico.

Remarks.--A female from Cueva de los Cuarteles is perhaps

also referable to this species. It is similar in all respects to

S^. lukensi except that it has no evidence of the lateral spermathecae.

The median spermathecae are very similar to those of S^, lukensi from

the type locality,

Schizomus davisi Gertsch, 1940

(Figs, 157, 177, 211; Tables 9, 11; Map 5)

Schizomus davisi Gertsch, 1940:1-4; Rowland, 1971c:117; Rowland,

1973a:21; Rowland, 1973c:135; Brignoli, 1974:149,

Holotype,--An adult male, taken at San Fernando, Tamaulipas,

Mexico, on 28 March 1937, by L. Irby Davis (AMNH, examined).

Description,--Male, Color brownish. Carapace with three

pairs of dorsal and two apical setae, Eyespots indistinct. Anterior

sternum with 13 bifid setae. Abdominal terga I-VII with two setae,

terga VIII-IX with four setae, segment XII without evidence of 177 posterodorsal process. Vestigial stigmata darker than sterna,

Flagellum spade shaped, with a pair of deep depressions undercutting a dorsal medially produced ridge, Pedipalpal trochanter produced apically into a tubercle which bears an apical seta, tarsal-basitarsal spurs about 1/5, claw about 1/2 length of tarsus-basitarsus,

Tarsal-basitarsal segments of leg I of the following approximate proportions: 28-6-7-6-8-9-19, Other leg segment measurements given

in Table 11,

Female unknown.

Comparisons,--S^. davisx is most similar to S^. mexicanus in form of the male's flagellum. The distal pits are, however, better defined in S^. davisi, The pedipalps are shorter and thicker than those of ^. mexicanus and the trochanter has a characteristic terminal

spine. S^. mulaiki is also closely related, but the males are readily distinguished by the much smaller, single distal depression.

Distribution.--This species is known only from a single

specimen from San Fernando, Tamaulipas, Mexico.

Remarks.--This species is probably a relict of a once widely

distributed conmon ancestor of S^, mexicanus, ^. reddel 1 i, S_. lukensi, i. bartolo, and _S. mulaiki. It has been searched for at the type

locality, but has not been rediscovered. It is most probable that

this specimen was collected near the banks of the Rio San Fernando,

since other areas surrounding San Fernando seem quite xeric and thus

quite unsuited for schizomids. Perhaps long term irrigational 178 drainage and drying of Rio San Fernando have eliminated S^, davisi from this part of its range.

Schizomus reddelIi Rowland, 1971

(Figs. 159, 175, 189, 190; Tables 9, 11; Map 5)

Schizomus reddelIi Rowland, 1971c:123, 124, 125, 126; Rowland, 1973a:

21; Rowland, 1973c:135; Brignoli, 1974:149,

Holotype.--An adult male, taken in Cueva de Tres Manantiales,

8 km NNE Chamal, Tamaulipas, Mexico, on 27 May 1968, by J. Reddell

(AMNH, examined).

Female.--An adult female, taken in Cueva de Tres Manantiales, in January 1972, by W. Russell (TTU),

Description.—Male. Color brownish. Carapace with two pairs of dorsal and two apical setae. Eyespots absent or only very indistinct. Anterior sternum with 10 bifid setae. Abdominal terga

I-VII with two setae, terga VIII-IX with four setae, segment XII without evidence of posterodorsal process. Vestigial stigmata darker than sterna. Flagellum lanceolate, with faint depressions medially, nearly flat dorsally. Pedipalpal trochanter produced slightly distally, tarsal-basitarsal spurs about 1/5, claw about 1/2 length of tarsus-basitarsus. Tarsal-basitarsal segments of leg I of the following approximate proportions: 37-7-9-8-10-10-20. Other leg segment measurements given in Table 11.

Female. Flagellum composed of three articles. Lateral spermathecae missing, medians long, thin, and slightly curved outward. 179

Comparisons,--S^, reddel Ii is generally larger than S^, mexicanus and the females differ in that S^, reddel Ii has a single pair of spermathecae, while S^, mexicanus has both. While the morphology of the male's flagellum in S^, mexicanus is, in most cases, distinct from S^. reddel 1 i, a variant from near Gomez Farias shows striking similarity to and is to me inseparable from that of S^, reddelli. The pedipalps of the males are sometimes dimorphic in S_, mexicanus, but are not so in S^, reddel Ii, See also under S^, mexicanus.

Distribution,--This species is known from Cueva de Tres

Manantiales and Cueva de los Vampiros, near Chamal, Tamaulipas, Mexico.

Remarks,--I have decided to maintain these populations as specifically distinct from that of its close relative, S^, mexicanus.

While the form of the male's flagella are sometimes inseparable, normally the best character in species recognition, the female's spermathecae have been found to be diagnostic, S^, reddel Ii probably represents a high altitude relict of a formerly widely ranging population ancestral to it and S^. mexicanus. S^. reddel Ii is probably troglobitic.

Additional records,--Tamaulipas: Cueva de Tres Manantiales,

January 1972, W, Russell, one female, one immature (TTU); Cueva de los Vampiros, 6 mi, NNE Chamal, 27 May 1968, J, Reddell, one male, three females, five immatures (TTU). 180

Schizomus mexicanus Rowland, 1971

(Figs. 2, 155, 156, 171, 172, 185-187, 207-210;

Tables 9, 11; Map 5)

Schizomus mexicanus Rowland, 1971c:117-119, 124, 125; Rowland,

1973a:10, 21, 22; Rowland, 1973b:200-201; Brignoli, 1973:6;

Rowland, 1973c:135, 137; Brignoli, 1974:141, 146-147, 151,

Holotype.--An adult male, taken in Sotano de la Tinaja, 10 km

NNE Ciudad Valles, San Luis Potosi, Mexico, on 18 February 1970, by

J. A. L, Cooke (AMNH, examined).

Allotype.--An adult female, taken with the holotype (AMNH, examined),

Paratypes,--An adult male and adult female, taken with the holotype (AMNH, examined).

Description.--Male. Color brownish. Carapace with two pairs of dorsal and two apical setae, Eyespots indistinct. Anterior sternum with 10 bifid setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segment XII with no evidence of posterodorsal process. Vestigial stigmata darker than sterna.

Flagellum ovoid, with a pair of subdistal pits. Pedipalpal trochanter produced apically slightly, other segments elongate, the tibia with a jutting spur apposible to the tarsus-basitarsus, tarsal-basitarsal spurs about 1/5, claw about 1/3 length of tarsus-basitarsus.

Tarsal-basitarsal segments of leg I of the following approximate proportions: 45-8-10-9-10-10-27. Other leg segment measurements given in Table 11. 181

Female, Flagellum composed of three articles. Median and lateral spermathecae outwardly divergent, medians somewhat larger than laterals, slightly expanded and sclerotized apically.

Comparisons,--This species is most similar to S^, reddel Ii and i- davisi. The trochanter of the pedipalp of the male is greatly produced in the latter species, which serves to distinguish it from

S^. mexicanus. The male's flagella are similar, but the dorsal depressions are more distinct and proximal in S^, davisi, The female of S^, davisi is not known, S^, reddel 1 i has a much longer flagellum in the male than the typical mexicanus males, S^, reddel Ii also lacks definition of a median depression, but this is perhaps not separable from a Gomez Farias type which is very similar in this character to S^. reddel 1 i. The lateral spermathecae are missing in females of S^. reddel 1 i, but are present in S^, mexicanus.

Distribution,--S^, mexicanus occurs in epigean habitats and caves from various localities in Tamaulipas and San Luis Potosi,

Mexico,

Remarks,--This species is a troglophile, but is widespread in epigean localities. It occurs sympatrically with A, lucifer and S^, cookeX, both troglobites, in Sotano de la Tinaja, Another troglobitic species, S^, mitchelli, which is very closely related to S^, cookei, apparently excludes S^, mexicanus from Cueva de la Florida and Cueva del Pachon. S^. mexicanus occurs in epigean localities immediately outside the entrance to both caves. Why S^. mexicanus can co-exist with S^. cookei in Sotano de la Tinaja, but cannot co-exist with the 182 extremely similar S^, mitchelli in Cueva del Pachon and Cueva de la

Florida probably owes its answer not to grossly different adaptive strategies in the two situations, but rather to the nature of the caves. The former cave is an immense cavern which receives frequent and enormous input of materials via surface drainage, but the latter caves receive much less insurgence of materials. This may provide a very different form of energy input in the two situations.

Considerable input of leaf litter in Sotano de la Tinaja may create the necessary dimensions which allow S^. mexicanus and S^, cookei to co-exist.

Some cavernicolous populations of ^, mexicanus (those occurring) in Cueva de Taninul n, 1, Grutas de Quintero, Sotano de Yerbaniz, and

Sotano de la Tinaja) have in common with S^, mitchelli and S^, cookei, a peculiar and otherwise unique extreme darkening of the vestigial stigmata. No explanation of this curious development is offered.

Variation,--There seems to be two forms of the male's flagellum, though intergrades may have escaped collection. Two males from the

Gomez Farias Roadcut have an elongated flagellum, which contrasts with the form from the type locality and found elsewhere within the range of the species. The pedipalps of the males are strongly sexually dimorphic in cave populations, but are more similar in epigean populations.

Additional records,--San Luis Potosi: Sotano del Arroyo, near

Los Sabinos, 25 November 1962, J, Reddell, one female, one immature

(AMNH); Cueva de Los Sabinos, 8 mi NE Valles, 27 January 1968, J, 183

Reddell, one male, three females, five immatures (TTU); Cueva de los

Monos, 14 km NE Ciudad Valles, 29 July 1970, J, Fish, one female (TTU);

El Sotano de las Piedras, 7,5 km NE Ciudad Valles, 28 March 1970, W,

Elliott, two females, five immatures (TTU); Sotano de Pichijumo, 7 mi

NE Ciudad Valles, 1 June 1968, J, Reddell, 2 females, six immatures

(TTU); 26 January 1969, J, Reddell, T, Mollhagen, A, Smith, two males, one female, one immature (TTU); Cueva Pinta, 8 mi NE Valles, 31

January 1969, J, Reddell, W. Russell, one female (TTU); Ventana

Jabali, 10 km NW Tamuin, 26 March 1964, T. Raines, D. McKenzie, B.

Bell, one female (AMNH); Sotano del Tigre, 10 mi NE Valles, 1 February

1968, J. Reddell, R, Mitchell, three males, three females, four

immatures (TTU); Sotano de la Tinaja, 10,5 km NE Ciudad Valles,

6 June 1964, J, Reddell, D. McKenzie, L, Manire, one male, two

females (AMNH); 13 March 1969, J, Reddell, one male, five females,

eight immatures (TTU); 18 February 1970, J, A. L, Cooke, three males,

six females, six immatures (AMNH); 24 May 1971, M, Brownfield, one

female, two immatures (TTU); 16 March 1972, G, Campbell, one male,

two immatures (TTU); 16 March 1972, R, Mitchell, one female, one

immature (TTU); 16 March 1972, D, Kiser, 1 female, two immatures (TTU);

20 February 1973, W. Graham, two females, four immatures (TTU); 20

February 1973, T. Mollhagen, one immature (TTU); Cueva de Taninul #1,

13 km SE Ciudad Valles, 29 March 1970, W. Elliott, four females, three

immatures (TTU); 5 June 1967, R. Mitchell, two females, three immatures

(TTU); 27 March 1967, two males, four females, four inmatures (TTU);

Cueva Grande, 6 September 1967, J. Reddell, D, McKenzie, one female 184

(AMNH); Cueva Chica, 10 mi SE Valles, 30 January 1969, J, Reddell, one male, five females, six immatures (TTU); 31 January 1968, J,

Reddell, J, George, two males, three females (TTU); 5 June 1967, R,

Mitchell, two males, five females, five immatures (TTU); 19 May 1971,

W, Elliott, one female (TTU); Cueva de Taninul #4, 9 mi E Ciudad

Valles, 11 July 1969, S, and J, Peck, one female (TTU); Valles,

19 July 1956, W. Gertsch, three immatures (AMNH); 8 mi N Valles,

28 November 1964, J, Reddell, one male (AMNH). Tamaulipas: Cueva del Nacimiento del Rio Frio, 6 km S Gomez Farias, 15 February 1970,

R, Mitchell, one female (TTU); 16 February 1970, S, Wiley, one male,

one female, one immature (TTU); 15 March 1972, R, Mitchell, one

female (TTU); Arroyo del Nacimiento del Rio Frio, 16 February 1970, four females, four immatures (TTU); Sotano de Jineo, 1 km NW Gomez

Farias, 17 May 1971, W. Elliott, one female (TTU); surface, near

Cueva de la Florida, 16 km SSW Mante, 20 July 1973, J, Rowland, J.

Reddell, three males, one immature (TTU),

Schizomus pallidus Rowland, 1975

(Figs, 166, 179, 196; Tables 9, 10; Map 5)

Schizomus pallidus Rowland, 1975:13-15, 17,

Holotype,—An adult male, taken in Cueva de Macinga, Tlilapan,

Veracruz, Mexico, on 5 March 1973, by J, Reddell (AMNH, examined).

Allotype.--An adult female, taken with the holotype (AMNH,

exami ned). 185

Paratypes,--An adult male, two adult females, and two immatures, taken with the holotype (AMNH, examined).

Description,--Male. Color pale brownish. Carapace with three pairs of dorsal, the medians very small, and two apical setae.

Eyespots indistinct. Anterior sternum with 11 bifid setae. Abdominal terga I-VII with two setae, terga VII-IX with four setae, segment XII with no evidence of posterodorsal process. Vestigial stigmata darker

than sterna, Flagellum lanceolate, with a pair of median depressions,

Pedipalpal trochanter produced distally, tarsal-basitarsal spurs about

1/5, claw about 1/2 length of tarsus-basitarsus, Tarsal-basitarsal

segments of leg I of the following approximate proportions:

56-10-12-10-11-12-26, Other leg segment measurements given in Table

10.

Female, Flagellum composed of three articles. Median and

lateral spermathecae divergent, the medians about twice as long as

laterals; both pairs evenly sclerotized along length.

Comparisons.--See under S^. portoricensis.

Distribution,--This species is known only from Cueva de

Macinga, Veracruz, Mexico.

Remarks.--The large size of this species is quite remarkable

within the mexicanus group. The eyespots of this species are

indistinct, but are nonetheless present. This is the only character

which would contraindicate that this species is troglobitic. The

very light pigmentation of the body leads me to believe that this 186 species may be undergoing regressive changes to the cave environment, but may have not yet lost the eyespots.

Schizomus portoricensis (Chamberlin), 1922

(Figs. 161, 167-170, 181, 199-206; Tables 9, 10a; Map 5)

Stenochrus portoricensis Chamberlin, 1922:11-21; Mello-Leitao, 1931:

19; Giltay, 1935:8; Werner, 1935:469; Takashima, 1943:93;

Rowland, 1973b:195, 197, 200; Brignoli, 1974:145.

Schizomus antilus Hilton, 1933:91-92; Giltay, 1935:6; Takashima,

1943:94. SUGGESTED SYNONYMY, Female types from Corall

Nuevo (1500 ft,), and near Havana, Cuba, reportedly deposited

in the Pomona College (California) collection, not examined.

Schizomus cavernicolens Chamberlin and Ivie, 1938:102, 103; Gertsch,

1940:4; Takashima, 1943:94; Rowland, 1971b:117; Brignoli,

1974:149. NEW SYNONYMY. Female holotype from Xkyc Cave

(=Actun Xkyc), Yucatan, Mexico, deposited in the American

Museum of Natural History, examined,

Schizomus longimanus Rowland, 1971b:119-120, 124, 125; Rowland,

1973a:13, 16, 22; Rowland, 1973b:197, 200; Rowland, 1973c:

135, 137; Brignoli, 1973:6, 7, 8, 9; Brignoli, 1974:143, 144,

146, 149, 151. NEW SYNONYMY. Male holotype and female

allotype from Cueva Cerro Hueco, 3 km SE Tuxtla Gutierrez,

Chiapas, Mexico, deposited in the American Museum of Natural

History, examined.

Schjzomu^s floridanus Muma, 1967:18-20; Rowland, 1971a:304. NEW

SYNONYMY. Female holotype from Ross and Castellow Han^.i-ock, 187

Dade County, Florida, United States, deposited in the American

Museum of Natural History, examined.

Holotype.--An adult female taken in November, 1899, at Coamo

Springs, Puerto Rico, and deposited in the Museum of Comparative

Zoology (MCZ), examined.

Paratypes.--Several females taken with the holotype, and also deposited in the Museum of Comparative Zoology, examined.

Description of a male and female from 1 km S Muna, Yucatan,

Mexico,--Male, Color brownish green. Carapace with two pairs of dorsal and two apical setae, Eyespots distinct, vaguely triangular.

Anterior sternum with nine bifid setae, posterior sternum with bifid setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segment XII with no evidence of posterodorsal process.

Vestigial stigmata slightly darker than sterna. Flagellum ovoid, with a pair of median depressions, Pedipalpal trochanter produced distally, other segments slightly elongate, Tarsal-basitarsal spurs about 1/7, claw about 1/3 length of tarsus-basitarsus, Tarsal- basitarsal segments of leg I of the following approximate proportions:

49-6-8-8-8-8-18, Other leg segment measurements given in Table 10a,

Female. Females differ from the males in the following respects: flagellum with 3 sections, pedipalps not elongate, first legs noticably shorter, eyespots less distinct, color less greenish.

Spermathecae with median and lateral lobes outwardly divergent; medians heavily sclerotized along entire length; laterals much reduced and weakly sclerotized. 188

Comparisons,--S^, portoricensis is most similar and cladistically most proximal to S^, pallidus Rowland, 1975, The flagellum is similar in males of the two species, but is longer in S^, pallidus (,53) than in

S_. portoricensis (,42), The females are separable on the basis of the flagella also, which is longer in S^. pallidus (.45) than in S^, portoricensis (,30). The carapacal length in both sexes is about 1,4 and that of S^. portoricensis is about 1.25 at largest, and is usually about 1.05. S^. pallidus has three pairs of dorsal carapacal setae, whereas S^. portoricensis has two pairs.

Within its range in Mexico S^, portoricensis can always be distinguished by its two pairs of dorsal carapacal setae and small

size. The only other Mexican species with only two pairs of dorsal

setae, S^. pecki Rowland and OTU #7 of the pecki group, are large, over 1.5 carapacal length. Elsewhere, S^. portoricensis occurs with

two other species that also have two pairs of dorsal carapacal setae.

These species are S^. rowlandi, a troglobite, and S^, armasi, a close relative of the latter, both from Cuba. These species, however, are much larger with carapacal lengths of about 1,7, Females of S_, portoricensis can be distinguished from females of all other species

with two pairs of dorsal setae, members of the simonis and cubanicus

groups, within its range by the long flagella in the latter groups,

which are about ,40 or longer.

The fonn of the spermathecae, though relatively constant in

form may not, considering the variability of this character in other

species, be saioly distinguished from that of S.. silvino, S_, pallidus 189 and a species from Grutas de Atoyac, Veracruz, which is perhaps referable to S^, sbordonii. Only a single female of the latter species is available. The heavy sclerotization of the median spermathecal lobes allows their perception through the genital sternite, without dissection. Fig, 167 illustrates the spermatheca as it generally appears through the genital sternite. Though the spermatheca is only rarely visible without dissection in other species, it remains an expediant means of identifying S^, portoricensis with some dependability.

Variations,--This widespread species apparently has several geographical isolates, I have measured females from three widely separate localities and report the descriptive statistics in Table 10,

The sexual population from Cueva Cerro Hueco, Chiapas, differs from the widespread sexual individuals encountered at lower elevations in Chiapas and in the Yucatan Peninsula by the consistantly greater size. In particular, the males from Cueva Cerro

Hueco always have greatly elongate first legs. However, the discovery of a single male at Ciudad del Carmen, Campeche, and another from the entrance of Cenote (=Cueva) de Hoctun, Yucatan, with first legs and carapacal lengths only slightly shorter than those of males from

Cueva Cerro Hueco shows that these character states are not unique to the latter population. Females from different populations are not easily distinguished, even from widely separated localities. It will be noted in Table 10, however, that the measurements differ somewhat from locality to locality, A single classification analysis of 190 variance, for example, shows that there is a significant (p < ,025) added variance component among populations from Yucatan, Florida, and the Galapagos Islands for length of the patella of the first leg,

Gabriel's (1964) sum of squares simultaneous test procedure (SS-STP), however, shows that the Florida and Yucatan populations, when tested at a .05 critical level, form a homogeneous set in regard to this same character.

The morphology of the spermatheca is unusually consistant throughout the range of S^, portoricensis. No other species I have

studied shows this much consistance even at the same locality. The

lateral lobes, however, seem to be much larger in the Cueva Cerro

Hueco population.

Remarks,--Males of S^, portoricensis are known from Chiapas,

Campeche, and Yucatan, Mexico, Nicaragua, Guatemala, and Puerto Rico.

Well collected locations in which males are unknown are Florida,

Jamaica, and the Galapagos Islands, These populations are suspected

to represent, at least in part, parthenogenetic clones. Since evidence

of males from outside Nuclear Central America is limited to a single

specimen from Puerto Rico it is suspected that outlying populations

are generally parthenogenetic. But since no consistant morphological

difference between parthenogenetic and sexual females has been found,

it is not yet possible, short of rearing them, to certainly determine

the reproductive nature of the populations in question.

Collections which I suspect represent parthenogenetic clones

come from Yucatan, Florida, Bermuda, Cuba, St. Croix (Virgin Islands), 191

Puerto Rico, Jamaica, Ecuador, Colombia, and the Galapagos Islands.

An introduced population at the Cambridge University Botanical

Gardens reported by Cloudsley-Thompson (1949), represented only by females, is also referable to this species. Populations from Oaxaca and Veracruz are represented in collections by single females, hence are questionable in respect to sexuality,

James R, Reddell, who collected most of the Yucatan material, brought an interesting observation to my attention. In making.these collections he noticed a remarkable difference in the sex ratios of collections from epigean and cave situations (Map 2). Fifteen collections from surface localities have yielded 25 males, 9 females, and 13 immature individuals, while 44 collections from caves have yielded 6 males, 108 females, and 81 immature individuals. A contingency table analysis of the above data shows that there is a significant (p < .001) difference in the data sets. This striking paucity of males leads me to believe that most caves contain parthenogenetic clones. It is highly unlikely, considering the nonvagile nature of schizomids, that females seek out cave habitats and that males, born to a cave environment, seek epigean habitats.

It seems far more likely that some populations of S^. portoricensis are parthenogenetic.

The distribution of sexual populations in epigean habitats and parthenogenetic clones in caves raises interesting questions about the reproductive nature of this species which unfortunately cannot be adequately answered without some as yet unavailable information. 192

Some important questions to answer are: (1) are the parthenogenetic

clones, both on the Peninsula and elsewhere, of polyphyletic origin;

(2) are parthenogenetic females able to back cross to the sexual

populations given the opportunity; and (3) is there a behavioral

difference between males and females such that females are well

suited and males poorly suited to a cavernicolous existence, S_,

portoricensis would seem most likely to be facultatively

parthenogenetic and the various clones, both local and widespread,

to have polyphyletic origins, that is, females, in the absence of males, through some cytological mechanism, may reproduce asexually.

Females, arising parthenogenetically, may further be able to back

cross to the sexual population.

The parthenogenetic nature of cave populations may be due to

an extreme unsuitability of males to cavernicolous environments.

Males and females of most species, as in S^, portoricensis, possess

a sexual dimorphism which probably allows some degree of niche

separation. Perhaps the behavioral differences between the sexes

are so great in this species that, while the female is suited to

cavernicolous life, the male is less suited. This may be reflected

by the fact that males are generally a darker green than females,

and have better defined eyespots, adaptations which seemingly could

occur where males are more often exposed to light.

The appearance that most areas other than the Yucatan

Peninsula are occupied by parthenogenetic clones may be an artifact

of collecting, S^, portoricensis may simply be rarer in epigean 193 habitats, but a bias for cave collecting in the Antilles may be the reason for the apparent rarity of males from that area. The entire

Antilles are represented by 26 cave collections totalling one mature male and 102 mature females, and 13 surface collections totally 24 mature females. The peninsula of Florida is represented, however, by 30 surface collections totalling 49 mature females and Santa Cruz

Island, Galapagos Islands, is represented by two surface collections totalling 25 mature females. Moreover, the same segregation of parthenogenetic and sexual populations may obtain at the cave entrance in the Antilles as well as in Yucatan, The relatively large surface collections from Florida and the Galapagos, hov/ever, suggest that these populations may be totally parthenogenetic.

Males may arise in outlying populations by dispersal from sexual populations, or may arise through some genetic mechanism from a parthenogenetic parent. Apparently the commonest means of maintaining thelytokous parthenogenicity is by reunion of haploid meiotic nuclei in pairs (White, 1970), in which case males may occasionally be produced; however, these males are usually rare and sterile. Such a paucity of information is available on the distribution of males in outlying areas that determination of its role is impossible. Planned cytological studies on individuals from parthenogenetic clones in Florida and

Yucatan, and individuals from sexual populations in Yucatan may give concrete evidence of the relationships of the sexual and asexual populations and may further clarify the status of males in outlying areas. 194

Distribution,--S^, portoricensis is known from Bermuda, Florida,

Campeche, Chiapas, Oaxaca, Veracruz, Yucatan, Belize, Guatemala,

Nicaragua, Cuba, Dominica, Jamaica, Puerto Rico, Virgin Islands,

Colombia, Ecuador, Galapagos Islands, and England (introduced). The

ancestral populations of S^, portoricensis probably arose in Nuclear

Central America sometime before the appearance of the greatest part

of the Yucatan Peninsula (Late Pliocene), After emergence of the

Yucatan shelf its ancestors, probably along with progenitors of S^,

goodnightorum, moved northward into the Peninsula, up to and then

beyond the Isthmus of Tehuantepec, and west of the Chiapas Highlands

to the Chiapa Depression, Once established and widespread on the

Peninsula the sexual species probably began giving rise to

parthenogenetic clones. From Yucatan the parthenogenetic females

must have rafted onto the Greater Antillean Islands and Florida,

From Florida or perhaps directly from Yucatan they have also reached

Bermuda. Overland dispersal to the south must have occurred more

slowly, but dispersal through Central America probably was accomplished

in the Pliocene when many faunal elements were dispersing toward South

America. Arrival and proliferation in South America probably antedated

the ultimate arrival and colonization of the Galapagos Islands. See

Map 5 for its generalized distribution.

Additional records.--Bermuda, Pembroke, 24 July 1915, P- H,

Pope, from stomach contents of toad, 1 female (MCZ),

United States, Florida: Dade County: Ross and Castellow

Hammock, 4 January 1958, H, V. Weems, Jr., under bark, 1 female (AMNH); 195

Station No. 1, 28 January 1959, R. E. Woodruff, 2 females, 8 immatures

(AMNH); Matheson Hammock, 12 September 1959, H. V, Weems, Jr,, under bark, 5 females, 2 immatures (AMNH); Matheson Hammock, 29 August 1961,

H. V, Weems, Jr., under bark, 1 immature, in leaf mold, 1 female

(AMNH); Matheson Hammock, 12 August 1959, R. E. Woodruff, 4 females,

4 immatures (AMNH); Matheson Hammock, 12 August 1959, H. A. Denmark,

12 females, 6 inmatures (AMNH); Royal Palm State Park, 27-29 December

1940, A. F. Archer, 1 female (AMNH); Miami Brickell Hammock, 30

November 1961, F, W, Mead, 1 immature (AMNH); Miami, 1 June 1940,

E, M, Miller, rotten wood, 2 inmatures (AMNH); Miami, 10 February

1965, R, E, Woodruff, 1 female (AMNH); 2-5 mi S Florida City, 1 April

1957, R, Forster, W, J, Gertsch, 14 females, 6 immatures (AMNH);

Everglades National Park, W81, N25, 28 January 1959, H, A, Denmark,

3 imuatures (AMNH); Everglades National Park, 11 March 1968,

Chickering, 1 immature (MCZ); Everglades National Park, 1 March 1952,

M. H. Muma, under stones, 4 immatures (AMNH); Everglades W Homestead,

W81.35, N25.20, 17 December 1962, W. Ivie, 1 immature (AMNH);

Homestead, W Mowry St., avocado groves, March 1968, 1 female (AMNH).

Monroe County: No Name Key, hardwood litter, Ber 222, 5 August 1972,

S. Peck, 1 female, 1 immature (MCZ); Upper Matacumbe Key, Islamadora,

Ber 225, 8 August 1971, S. Peck, 1 female, 3 immatures (MCZ);

Everglades National Park, Gumbo Limbo Trail, 8 April 1969, No. 480 litter, C. Alteri, 3 immatures (AMNH); Key Vaca, Marathon, palm-hardwood litter, Ber 223, 7 August 1972, S. B. Peck (MCZ);

Stock Island, 12 May 1961, H. V. Weems, Jr., under rock in hammock, 196

1 female (AMNH); Marathon, 15 March 1968, S, Peck, Ber 111, litter and soil in scrubforest solution pocket, 1 female, 4 immatures (MCZ);

Marathon, 18 March 1968, S. Peck, palm litter, 7 immatures (MCZ);

Marathon, 15 March 1968, scrubforest, 2 females, 3 immatures (AMNH);

Lignum Vitae Key, Islamorada, 14 March 1968, S, Peck, B109, hardwood scrub litter, 2 immatures (MCZ); Key Largo Key, 30 January 1959,

H. V, Weems, Jr,, under rock, 3 immatures (AMNH); Little Duck Key,

W81.15, N24.40, 28 December 1963, J. and W. Ivie, 1 female (AMNH).

St. Lucie County: Ft. Pierce, rotten limb of Brazilian Pepper,

1 January 1965, M. H. Muma, H. L. G., 1 immature (AMNH).

Mexico. Campeche: Ciudad del Carmen, 28 July 1948, C. J. and M. L. Goodnight, 1 male (AMNH); Cenote de Cantemo, 1 km N

Cantemo, 18 December 1974, J. Reddell, D, McKenzie, L, Elliott,

4 females, 3 immatures (TTU); Actun Halmensura, 5 km E Cumpich,

31 October 1974, J, Reddell, D, McKenzie, S. Wiley, 1 female,

3 immatures (TTU); 10 km N Hopelchen, 27 July 1973, J, M, Rowland,

J. R. Reddell, 2 males, 4 immatures (TTU); 5 km SSW Ich-Ek, 27 July

1973, J. M. Rowland, J. R. Reddell, 1 male (TTU); Grutas de San

Antonio, 10 km ENE Bolonchenticul, 3 November 1974, J. Reddell, D.

McKenzie, S. Wiley, 3 females, 3 immatures (TTU); Grutas de San

Antonio, 23-24 November 1974, J. Reddell, D. McKenzie, S. Wiley,

12 females, 10 immatures (TTU); Grutas de Xtacumbilxunam, 5 km SW

Bolonchenticul, 29-30 July 1973, R. W. Mitchell, 1 female (TTU);

Grutas de Xtacumbilxunam, 19 April 1973, J. Reddell, D. McKenzie,

M. McKenzie, S. Murphy, M. Butterwick, 5 females, 3 immatures (TTU); 197

Grutas de Xtacumbilxunam, 25 November 1974, J, Reddell, D, McKenzie,

S. Wiley, 1 female (TTU), Chiapas: Cacahoatan, 10 mi NE Tapachula,

8 September 1950, C, J, and M, L. Goodnight, 1 female, 1 immature

(AMNH); Cueva Cerro Hueco, 3 km SE Tuxtla Gutierrez, 18 August 1967,

J. Reddell, J. Fish, M. Tandy, 3 males, 4 females, 7 immatures (AMNH);

Hoyo de Don Nicho, 8 mi W Ocozocoautla, 16 August 1967, J. Reddell,

J. Fish, T, Evans, 1 female (TTU); Ocosingo Valley, 1-7 July 1950,

C. J. and M. L. Goodnight and Stannard, 1 female (AMNH); 0.8 km N

Ruinas de Palenque, 25 July 1973, J. Reddell, R. W. Mitchell, 3 males,

5 females, 3 immatures (TTU); Ruinas de Palenque, 25 July 1973, J.

Reddell, J, M, Rowland, 2 males, 2 females, 5 immatures (TTU); El

Real, on hill 27 mi E Ocosingo, 3 July 1950, C. J. and M. L.

Goodnight and Stannard, 1 male, 3 females, 2 immatures (AMNH); between San Antonio and El Real, 9 July 1950, C, J, Goodnight and

Stannard, 2 females (AMNH); San Antonio, 12 mi E Ocosingo, 29-30

June 1950, C, J. and M. L, Goodnight and Stannard, 2 females, 2 immatures (AMNH); Cueva del Tempisque, 8 mi W Ocozocoautla, 17

August 1967, J, Reddell, T. R, Evans, 6 females, 8 immatures (TTU),

Oaxaca: 10 mi S Tomellin, 14 August 1967, J. Reddell, J, Fish, T,

Evans, 1 female (TTU), Veracruz: Penuela, W96,48, N18,53, 26 April

1963, W, J. Gertsch and W. Ivie, 1 female, 1 immature (AMNH).

Yucatan: Cenote de Acanceh, Acanceh, 8 October 1974, J. Reddell,

1 female (TTU); Cueva de Aguacate, 2 km S Maxcanu, 17 October 1974,

J. Reddell, R. Solis, D. McKenzie, S. Wiley, 2 females (TTU); Grutas de Balankanche (=Cueva Bolonchen), 4 km E Chichen Itza, July 1948, 198

C. J. Goodnight, 1 female (AMNH); Grutas de Balankanche, 10-12

December 1974, J, Reddell, D, McKenzie, S, Wiley, 2 females,

3 immatures (TTU); Becanchen, 1 August 1973, J, Reddell, 1 male,

1 immature (TTU); Actun Chacaljas, 3 km S Calcehtok, entrance sink,

3 August 1973, J, Reddell, 1 male (TTU); Actun Chunup, 2 km E

Maxcanu, 16 October 1974, D, McKenzie, S, Wiley, 1 female (TTU);

Dry Cenote (=Cenote Seco), Chichen Itza, 10 July 1948, C. J.

Goodnight, 1 male, 1 female (AMNH); Actun Gongora (=Gongora Cave),

Oxkutzcab, lots 109 and 111, A. S. Pearse, 2 females, 1 immature

(AMNH); Cenote (=Cueva) de Hoctun, Hoctun, 16 March 1973, J. Reddell,

D. McKenzie, M. McKenzie, S. Murphy, M. Butterwick, 2 males, 1 female, 1 immature (TTU); Cenote (=Cueva) de Hoctun, entrance area,

12 August 1973, J. Reddell, 1 male (TTU); Hoctun, 12 August 1973,

J. Reddell, 1 male (TTU); Cenote Hunto Chac (Cueva del Pozo), 12

April 1973, J. Reddell, M. McKenzie, 1 female (TTU); Tres Linteles,

Chichen Itza, 6 July 1948, C. J, Goodnight, 2 males (AMNH); Chichen

Itza, C, J, Goodnight, 1 female (AMNH); Chichen Itza, 8 August 1973,

J. Reddell, 2 males, 3 females, 2 immatures (TTU); Pyramid, Izamal,

10 August 1973, J, Reddell, 1 male, 1 female, 1 immature (TTU);

Actun (=Cueva de) Kaua, 1 km S Kaua, 8 January 1972, D, McKenzie,

2 females (TTU); Actun Kaua, 9-10 October 1974, J, Reddell, D,

McKenzie, S, Wiley, 1 female (TTU); Cenote Kabahchen, Mani, 5 October

1974, J. Reddell, D. McKenzie, S. Wiley, 2 females, 2 immatures (TTU);

Actun Kiuick, Kiuick, 13 November 1974, J. Reddell, D. McKenzie, S.

Wiley, 1 female, 1 immature (TTU); Grutas de Loltun, Oxkutzcab, 1 199

January 1972, D. McKenzie, 1 male, 1 female (TTU); Cenote (=Cueva)

Luchil, 3 km S Merida, October 1974, J. Reddell, S. Wiley, 1 immature

(TTU); Ruinas de Mayapan, 14 August 1973, J. Reddell, 4 males, 1 female, 1 immature (TTU); 1 km S Muna, 31 July - 4 August 1973, J.

Reddell, 2 males, 1 female, 2 immatures (TTU); Cueva de Orizaba,

Orizaba, 6 km S Buena Ventura, 1 April 1973, J. Reddell, D, McKenzie,

M, McKenzie, S, Murphy, M, Butterwick, 2 females, 3 immatures (TTU);

Cenote de Orizaba, Orizaba, 6 km S Buena Ventura, 1 August 1973,

J. Reddell, S. Murphy, 1 immature (TTU); Oxkutzcab, 31 July 1973,

J. Reddell, 2 immatures (TTU); 7 km S Oxkutzcab, berlese, 31 July

1973, R, W, Mitchell, J, R, Reddell, 1 male (TTU); Cenote de Paca,

7 km E Tikuch, 11 April 1973, S, Murphy, 1 female, 1 immature (TTU);

Actun Sabaca, 6 km S Tekax, 4 December 1974, J, Reddell, D, McKenzie,

S. Wiley, J. Andrews, R. W, Mitchell, 4 females, 3 immatures (TTU);

Cenote de Sambula, Kopoma, 15 October 1974, J. Reddell, D. McKenzie,

S. Wiley, 1 female (TTU); Cenote de Sambula, Motui, 28 March 1973,

J. Reddell, S. Murphy, 5 females, 3 immatures (TTU); Cueva de San

Isidro, Merida, 21 March 1973, J. Reddell, S, Murphy, 1 immature

(TTU); Cenote de San Jose, Merida, 6 October 1974, J, Reddell, D,

McKenzie, 1 female (TTU); Cenote de Sihunchen, Sihunchen, 23 March

1973, J, Reddell, M, McKenzie, S, Murphy, 1 female, 3 immatures

(TTU); Cueva Sodzil, 5 km W Sucopo, 31 March 1973, J, Reddell, D.

McKenzie, M, McKenzie, S. Murphy, 4 females (TTU); Cenote Sucopo,

Sucopo, 13 March 1973, J, Reddell, S, Murphy, 1 female, 5 immatures

(TTU); Cueva de Tecoh, Merida, 6 October 1974, J, Reddell, D. 200

McKenzie, 1 immature (TTU); Cueva de Tecoh, October 1974, J, Reddell,

S. Wiley, 1 female, 1 immature (TTU); Tixcocob, 12 August 1973, J.

Reddell, 5 males (TTU); Grutas de Tzab-Nah, 2 km S Tecoh, 26 April

1973, J. Reddell, M. McKenzie, 5 females, 2 immatures (TTU); Grutas de Tzab-Nah, 22 April 1973, J. Reddell, D. McKenzie, M, McKenzie,

1 female, 1 immature (TTU); Grutas de Tzab-Nah, 10 October 1974, J,

Reddell, D. McKenzie, S. Wiley, 11 females, 14 immatures (TTU);

Actun Xkyc (=Xkyc Cave), Calcehtok, 6 August 1936, A. S, Pearse,

1 female (AMNH); Actun Xpukil, 4-5 April 1973, J. Reddell, D.

McKenzie, M, McKenzie, S, Murphy, 1 female (TTU); Actun Xpukil,

18-19 March 1973, J, Reddell, S, Murphy, D, McKenzie, M, McKenzie,

M. Butterwick, 3 females, 4 immatures (TTU); Actun Xpukil, entrance sink, 3 August 1973, J, Reddell, 10 females, 9 immatures (TTU);

Actun Xpukil, 18-19 March 1973, J, Reddell, D. McKenzie, M. McKenzie,

S. Murphy, M. Butterwick, 4 females, 7 immatures (TTU); Actun Ziz,

Oxkutzcab, 3 December 1973, J. Reddell, A, Gamboa, D, McKenzie, S,

Wiley, R. W. Mitchell, 1 male, 1 immature (TTU),

Belize (=British Honduras). Caves Branch, 4-14 August 1972,

S. and J. Peck, 1 immature (AMNH); Augustine, 1500 ft, Rio Frio Cave

B, 30 July 1972, 20 August 1972, S. and J. Peck, 1 female, 1 immature

(AMNH); Augustine, 1500 ft, Rio Frio Cave A, 20 July 1972, S, Peck,

1 female, 1 immature (AMNH); Cave near Augustine, July 1972, C.

Goodnight, 1 female, 1 immature (AMNH); near Augustine, Rio Frio,

20 July 1972, C, and M, Goodnight, 1 female (AMNH); Hummingbird

Highway, 2 August 1972, C. J. Goodnight, 1 female (AMNH); Caves 201

Branch, Buck's Bypass Cave, 14 August 1972, S, Peck, 1 female;

Belmopan, 13 August 1972, S, and J. Peck, 1 female (AMNH); Belmopan,

7 August 1972, S, Peck, 1 female, several eggs (AMNH),

Guatemala, Alta Verapaz: Near village of Lanquin, June

1959, W, W, Varnedo, 2 females (AMNH); Grutas de Lanquin, Lanquin,

28 August 1969, S, and J, Peck, 10 females (AMNH), El Peten:

Najohnaj Coholtunich, 25 August 1972, S, and J, Peck, 4 males, 8 females, 6 immatures (AMNH).

Nicaragua. Musawas, 1-4 November 1955, 1 female; Musawas,

Waspic River, 10 October 1957, B, Malkin, 1 female, 1 immature (AMNH).

Cuba, S. A, Banos, Habana, 11 August 1971, L, F, deArmas,

1 female, 2 immatures; Cienfuegos, central Soledad, 2 July 1957, 1 female (LFA); Cueva Bellamar, Matanzar, 800 ft from entrance, January

1958, 1 female (AMNH),

Dominica. Roseau, botanical gardens, 2 February 1968, B,

Malkin, 7 females (AMNH); Salibia, under rotten banana leaves, 28-29

January 1968, B, Malkin, 1 female (AMNH),

Jamaica, St, Andrews Ferry, 0,9 mi W Red Hills, 1 female

(MCZ). St. Ann Parish: Falling Cave, Douglas Castle, 3 November 1973,

R. Norton, 1 female (AMNH); Hutchinson Hole Cave, 27 March 1973, R.

Norton, R. Zimmerman, 1 female (AMNH); "Rambriber" Cave, Douglas

Castle, 20 December 1972, S. and J. Peck, 1 female (AMNH); St, Catherine

Parish: Guanaboa, 28 February 1957, Chickering, 1 female (MCZ); 1,5 mi

SW Ewarton, 1000 ft, 7 April 1968, S, Peck, A, Fisher, 2 females, 4 immatures (AMNH); St. Claire Cave, Ewarton, 27 December 1972, S. and 202

J. Peck, 1 female (AMNH); Guanaboa, 4 December 1957, Chickering, 2 females (MCZ), St. Elizabeth Parish: Wallingford Caves, Wallingford,

27 August 1974, S, Peck, 3 females (MCZ), St, James Parish: Maldon

School Cave, Maldon, 2 September 1974, S, Peck, 1 female, 2 immatures

(MCZ); Brandon Hill Cave, Montego Bay, 1 September 1974, S, Peck, 3 females, 7 immatures (MCZ), Manchester Parish: Oxford Cave,

Auchtembeddie, 27 August 1974, S. Peck, 1 female, 5 immatures (AMNH).

St. Mary Parish: Luck Hill Cave, 31 March 1973, R. Zimmerman, 2 females (MCZ). Trelawny Parish: Dormilly Cave, 2 mi NE Deeside, 28

August 1974, S. Peck, 14 females, 8 inmatures (AMNH); Deeside Cave,

Deeside, 28 August 1974, S, Peck, 10 females, 3 immatures (MCZ);

Printed Circuit Cave, 30 March 1973, R, Norton, R, Zimmerman, 3 females, 3 immatures (AMNH); Carambie Cave, Spring Garden, 4 September

1974, S. Peck, 14 females, 3 immatures (MCZ); Harties Caves, Spring

Garden, 4 September 1974, S, Peck, 2 females, 2 immatures (MCZ),

Westmorland Parish: Roaring River Cave, 7 mi NE Savannala Mar, 29

August 1974, S, Peck, 7 females, 11 immatures (AMNH),

Puerto Rico, Aguas Buenas Cave, Aguas Buenas, 30 December 1966,

S. Peck, 2 females, 2 inmatures (AMNH); Aguas Buenas Cave, Ent.

Caguitas, 0-500 ft, 14 May 1973, S, Peck, 1 female (MCZ); Aguas Buenas

Cave, near Hoyo Petroglyph, 250 m, 15 May 1973, S, Peck, 1 female

(AMNH); Aguas Buenas Cave, Aguebana, 250 m, 7-17 May 1973, S. Peck,

1 female (MCZ); Cueva de los Alferos, Barrio Moza, near Isabela, 29

December 1966, S. Peck, 2 females (AMNH); Cueva Los Chorros, 15 km S

Arecibo, 31 May 1974, S. and J. Peck, 10 females, 4 immatures (MCZ); 203

Coamo Springs, 1899, several females (MCZ); Coamo Springs, 24 March

1906, W. M. Wheeler, 5 females (MCZ); Cueva El Convento, Guayanilla,

12 June 1974, S. and J. Peck, 12 females, 11 immatures (AMNH); Cueva

El Convento, guano berlese, 12 June 1974, S, Peck, 7 females, 15 immatures (MCZ); Cueva de Corozal, Corozal, 4 January 1967, S. Peck,

1 female (AMNH); Guajataca Forest near Publito de Ponce, 4 July 1958,

Sanderson, 1 female (MCZ); Mayaquez, 19 January 1964, Chickering, 1 female (MCZ); Mayaquez, 16 January 1964, Chickering, 1 female (MCZ);

Cueva Tuna, 3,5 km S Cabo Rojo, 13 June 1974, S, and J, Peck, 1 male,

3 females, 1 immature (AMNH).

Virgin Islands. St. Croix: Christiansted, 1940, H. A. Beatty,

1 female (MCZ),

Colombia, Near Cali, 2 October 1969, Eberhardt, 1 female

(MCZ).

Ecuador. Guayaquil, 31 October 1942, 2 females (AMNH);

Galapagos Islands, Isla Santa Cruz, Horneman Ranch, 16 February 1964,

D. Q. Cavagnaro, R. 0. Schuster, 3 females, 1 immature (CAS); Santa

Cruz Island (Indefatigable), November 1964, J. and N, Leleup, 22 females, 6 immatures (ISB),

England. Cambridge, University Botanical Gardens,

Cloudsley-Thompson, 3 females (AMNH). Table 10.--Descriptive statistics for 12 females of S^. portoricensis from each of the following localities: 1, Actun Xkyc,

Yucatan; 2, Matheson Hammock, Dade County, Florida; 3, Santa Cruz

Island, Galapagos Islands; and 4, nine males from Hoctun and Tixcocob,

Yucatan, SD, standard deviation, CV, coefficient of variation. 205

TABLE 10

1 2

Range Mean SD CV Range Mean SD CV

1.01-1.12 1,067 .036 3.37 1.00-1,08 1.048 .023 2,19

.25-.27 ,262 .008 3.05 ,24-,27 .255 .009 3,53

.92-1.10 .972 .054 5,56 ,89-1,00 .949 .036 3.79

1.01-1.26 1.109 ,069 6,22 .99-1,19 1.109 .050 4,51

.72-.84 .781 ,037 4,74 .69-,83 .773 .034 4,40

.68-.76 .711 .030 4,22 ,62-,74 .692 .036 5,20

.68-.78 .715 ,028 3,92 ,63-.74 .694 .030 4,32

.36-,44 .398 ,020 5,03 ,34-.41 .394 .021 5,33

.42-,50 ,451 ,028 6,21 .39-.46 .431 .024 5,57

.38-,44 ,406 ,022 5,42 .36-.42 .393 .019 4.83

,59-,70 ,645 .033 5,12 .56-.66 .618 .028 4.53

,26-.33 ,294 ,018 6,12 .26-.31 .289 ,020 6.92

.33-,39 .355 ,018 5,07 .30-.38 .338 ,025 7.40

,40-,46 ,434 .020 4,61 .40-.46 .428 ,020 4.67

,98-1.14 1.040 ,046 4,42 .94-1.06 1.012 .032 3,16

.40-.50 .454 ,027 5,95 .42-.48 .454 .018 3,96

.68-.83 .734 ,041 5.59 .68-.76 .728 .028 3,85

,52-.65 .600 ,036 6.00 .55-.62 .593 .021 3,54 206

TABLE 10 (conti nued)

3 4

n Range Mean SD CV Range Mean SD CV

1 1,00-1,10 1,048 .027 2.58 1,02-1,26 1.096 .081 7,37

2 ,25-,28 .266 .008 3,01 ,32-.36 .350 .015 4,29

3 - - - - ,21-.25 .24 .016 6,62

4 ,91-1,05 .986 .043 4,36 1,18-1,64 1.313 .167 12,70

5 1,06-1,22 1.148 .044 3,83 1.42-2,10 1.659 .237 14,28

6 ,75-,87 .802 .033 4,11 1.04-1,48 1.205 ,177 14,69

7 ,69-,75 ,727 .019 2,61 ,80-,99 .895 ,067 7,48

8 ,65-.76 ,723 ,028 3,87 ,70-,92 .763 ,080 10.46

9 ,38-,43 ,408 ,016 3,92 ,41-,54 .456 ,054 11.84

10 ,41-,46 ,443 .018 4,06 ,44-,65 .518 ,080 15.40

11 ,38-,45 ,419 .021 5,01 ,42-,53 .456 ,042 9.22

12 .59-,68 .638 .034 5.33 ,61-,83 .676 .070 10.39

13 ,28-.33 .306 .015 4.90 .29-.39 .317 .034 10.71

14 .32-.39 .362 .021 5.80 ,33-,46 .371 .046 12,53

15 .41-.48 .444 .019 4.28 .42-.55 .460 ,041 8,96

16 .97-1.11 1,051 ,041 3,90 1.02-1.40 1.146 ,123 10,70

17 .42-.50 ,469 ,028 5,97 .45-.71 .546 ,078 14,25

18 .69-.78 ,747 ,029 3,88 .73-1.10 .858 ,124 4,48

19 ,57-,67 .631 ,027 4.28 .59-.73 .630 .049 7.78 207

Schizomus sp,, OTU #11

(Fig, 195; Tables 9, 11; Map 5)

Specimens examined.--An adult female and an immature, taken

6 mi S Valle Nacional, Oaxaca, Mexico, 2000 ft., on 19 May 1971, by

S. Peck (TTU),

Description,--Male unknown. Female, Color brownish green.

Carapace with three pairs of dorsal and two apical setae. Eyespots distinct. Anterior sternum with nine bifid setae. Abdominal terga

I-VII with two setae, terga VIII-IX with four setae. Vestigial stigmata darker than sterna, Flagellum composed of three sections.

Pedipalpal trochanter slightly produced distally, tarsal-basitarsal spurs about 1/5, claw about 1/2 length of tarsus-basitarsus,

Tarsal-basitarsal segments of leg I of the following approximate proportions: 28-4-5-5-5-6-14, Other leg segment measurements given in Table 11. Median spermathecae almost horizontally divergent, extremely long, lateral spermathecae short, dome shaped; medians terminate in poorly defined sclerotized bulbs.

Comparisons,--See under S^, moisii,

Distribution,--This taxon is known only from near Valle

Nacional, Oaxaca, Mexico.

Remarks.--See under S. moisii. 208

Schizomus moisii Rowland, 1973

(Figs. 163, 176, 197; Tables 9, 11; Map 5)

Schizomus moisii Rowland, 1973c:136, 137-139, 140; Rowland, 1975:13.

Holotype.--An adult male, taken in Grutas de Monteflor, 6 km

NE Valle Nacional, Oaxaca, Mexico, on 28 December 1972, by J. Reddell,

D. McKenzie, and S. Murphy (AMNH, examined).

Allotype.—An adult female, taken with the holotype (AMNH, examined).

Paratypes.--Five adult males, five adult females, and one juvenile, taken with the holotype (TTU, examined).

Description.--Male. Color brownish green. Carapace with three pairs of dorsal and two apical setae. Eyespots distinct.

Anterior sternum with nine bifid setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segment XII with no evidence of posterodorsal process. Vestigial stigmata darker than sterna. Flagellum diamond shaped, with a pair of median depressions flanked laterally by a pair of swellings. Pedipalpal trochanter produced distally, tarsal-basitarsal spurs about 1/5, claw about 1/2 length of tarsus-basitarsus. Tarsal-basitarsal segments of leg I of the following approximate proportions: 44-7-8-7-7-9-16. Other leg segment measurements given in Table 11.

Female. Flagellum composed of three sections. Spermathecae with medians three times longer, but somewhat thinner than laterals, the latter being dome shaped, medians terminate in a poorly defined sclerotized bulb. 209

Comparisons.--S^, moisii and its closest relative, OTU #11, are distinct from others of the mexicanus group in that they possess three pairs of dorsal carapacal setae, distinct eyespots, greenish coloration and short, wide lateral spermathecae. S_. moisii has straight, short, thick median spermathecae, whereas in OTU #11 the medians are apically outwardly divergent, very long, and relatively thin.

Distribution,--This species is known only from Grutas de

Monteflor, Valle Nacional, Oaxaca, Mexico,

Remarks,--While this species is known only from a cave, its dark pigmentation indicates that it is a facultative cave inhabitant and should be found in epigean situations in the area of the cave. It occurs with OTU #8 of the pecki group, which may, however, be a troglobite, A comparable situation obtains in Grutas del Cocona,

Tabasco, where the dark green S^, trilobatus occurs in the twilight zone and the troglobitic S^, pecki, another pecki group member, occurs further back.

The very close similarity and geographical proximity of S^, moisii and OTU #11 leaves some doubt that they are actually separate species, I have treated them separately here, however, because of the great disparity in the morphology of the females' spermathecae.

Variation.--The apex of the carapace of the holotype is typical of most other schizomids; however, two of the paratypes show variation in this character unlike any I have seen before. Two of the male paratypes have no conical process at the apex of the carapace, but merely a truncate margin, which is nearly straight from 210 the right side to the left. The two apical setae usually present are reduced to one, the distal seta probably being lost, I cannot speculate on the significance of this phenomenon.

Schizomus cookei Rowland, 1971

(Figs, 165, 180, 183, 184; Tables 9, 11; Map 5)

Schizomus cookei Rowland, 1971b:122-123, 124, 125; Rowland, 1973c:

135; Dumitresco, 1974:291; Brignoli, 1974:146,

Holotype,--An adult male, taken in El Sotano de la Tinaja,

10 km NNE Ciudad Valles, San Luis Potosi, Mexico, on 19 February 1970, by J. A, L, Cooke (AMNH, examined).

Allotype.--An adult female, taken with the holotype (AMNH, examined).

Paratypes,--An adult male and an adult female, taken with the holotype (AMNH, examined).

Description,--Male, Color brownish. Carapace with three pairs of dorsal and two apical setae, Eyespots absent. Anterior sternum with 11 bifid setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segment XII with no evidence of posterodorsal process. Vestigial stigmata very much darker than

sterna. Flagellum triangular, with a median pit flanked by lateral

swellings. Pedipalpal trochanter produced distally, other segments

elongate, the tibia with large spur apposible to tarsus-basitarsus,

the latter emarginate with spurs about 1/8, claw about 1/3 length of

tarsus-basitarsus. Tarsal-basitarsal segments of leg I of the 211 following approximate proportions: 35-6-8-7-9-10-22. Other leg segment measurements given in Table 11,

Female. Flagellum composed of three articles. Spermathecae composed of four or five lobes, the medians possibly trifurcate, the terminations not sclerotized, the medians the largest.

Comparisons,--See under S^, mitchelli.

Distribution.--This species is known from El Sotano de la

Tinaja and El Sotano de Yerbaniz, San Luis Potosi, Mexico,

Remarks.--This species occurs sympatrically with S^, mexicanus and A. lucifer, and is, like its closest relative S^. mitchelli, troglobitic. Communication between the above mentioned caves must be subterranean since epigean habitats are exclusively occupied by

S^. mexicanus and perhaps P^. pachypalpus. Also see under S^, mexicanus.

Additional records,--San Luis Potosi: El Sotano de Yerbaniz,

22,5 km N Ciudad Valles, 8 January 1971, W, Elliott, one female (TTU),

Schizomus mitchelli Rowland, 1971

(Figs, 164, 182, 188, 212-215; Tables 9, 11; Map 5)

Schizomus mitchelli Rowland. 1971b:121-122, 124, 125; Brignoli, 1973:6;

Rowland, 1973c:135; Brignoli, 1974:141, 145-146, 148, 151.

Holotype.--An adult male, taken in La Cueva del Pachon, 15 km

SSW Ciudad Mante, Tamaulipas, Mexico, on 25 November 1967, by S. Fowler

and J. Reddell (AMNH, examined).

Allotype.—An adult female, taken in La Cueva del Pachon, on

6 June 1967, by R. Mitchell (AMNH, examined). 212

Paratypes.--An adult male and an adult female, taken with the allotype (AMNH, examined).

Description,—Male, Color brownish. Carapace with three pairs of dorsal and two apical setae, Eyespots absent. Anterior sternum with 11 bifid setae. Abdominal terga I-VII with two setae, terga

VIII-IX with four setae, segment XII with no evidence of posterodorsal process. Vestigial stigmata very much darker than sterna, Flagellum triangular, with a median pit flanked by slight lateral elevations,

Pedipalpal trochanter produced distally, other segments elongate, tibia with large spur apposible to tarsus-basitarsus, the latter emarginate with spurs about 1/8, claw about 1/3 length of tarsus- basitarsus, Tarsal-basitarsal segments of leg I of the following approximate proportions: 35-6-6-7-8-7-19. Other leg segment measurements given in Table 11.

Female. Flagellum composed of three articles. Spermathecae composed of three or four lobes, the medians possibly trifurcate; the terminations not sclerotized, the laterals the largest.

Comparisons.--The closest relative of S^. mitchel Ii is S^. cookei. They are distinct from other species in the group in the possession of three pairs of dorsal carapacal setae, wide male flagella, a single median depression and strongly dimorphic pedipalps.

S. mitchelli has no elevations basolateral to the median flagellar pit in the males whereas S^, cookei males have them. The female's median spermathecal lobes are narrower than the laterals in S^,

IBltchelLL. but just the opposite in S^, cookei. 213

Distribution.--This species is known from La Cueva del

Pachon, Cueva de la Florida, and Grutas de Quintero, Tamaulipas,

Mexico.

Remarks.--This species, like its closest relative ^, cookei, is troglobitic. Any communication between the above mentioned caves must be subterranean, since epigean habitats are exclusively occupied by S^. mexicanus and perhaps _P, pachypalpus. Also see under

S^. mexicanus.

Additional records,--Tamaulipas: La Cueva del Pachon, 6

June 1967, R, Mitchell, one immature (TTU);8 June 1967, J, Reddell, one male (TTU); 25 November 1967, J, Reddell, two females (TTU); 12

March 1969, J. Reddell, one male, three females, five immatures

(TTU); 10 July 1969, S, and J, Peck, R, Norton, one male, four females, six immatures (TTU); Cueva de la Florida, 8 km NE Antiguo

Morelos, 15 km SW Ciudad Mante, 28 May 1968, J. Reddell, one male, five females, five immatures (TTU); 10 March 1969, J. Reddell, S,

Fowler, B, Cook, five males, six females, seven immatures (TTU); 16

February 1970, J. Cooke, two immatures (AMNH); 20 May 1971, J, Cooke,

M. Brownfield, one male, four females, four immatures (TTU); 19

February 1973, S. Wiley, one male (TTU); Grutas de Quintero, Quintero,

6 July 1969, S, Peck, three males, one female, one immature (TTU); 1

June 1972, W. Peck, one male, one female (MCZ), Figs. 155-163.--Dorsal views of male's flagella of the mexicanus group: 155, 156, S^, mexicanus: 155, from the type locality; 156, from Gomez Farias; 157, S^. davisi; 158, S^. bartolo; 159, S^. reddel Ii;

160, S^. lukensi; 161, S^. portoricensis from Cueva Cerro Hueco, Chiapas;

162, S, mulaiki; 163, S, moisii. Scale: 1 cm = .06 mm.

Figs, 164-170.—Parts of schizomids of the mexicanus group:

164-166, dorsal views of male's flagella: 164, S^, mitchelli; 165,

S_. cookei; 166, S^. pallidus; 167-170, female S^. portoricensis: 167, ventral view of abdominal sterna II and III, showing spermathecae through the integument; 168, dorsal view, legs and pedipalps past the trochanter omitted; 169, lateral view of flagellum; 170, lateral view of right pedipalp. Scale: Figs, 164-166, 169, 170, 1 cm =

,06 mm; Fig. 167, 1 cm = ,2 mm; Fig, 168, 1 cm = ,4 mm.

Figs. 171-182,--Lateral views of male's flagella of the mexicanus group: 171, 172, S^, mexicanus: 171, from the type locality; 172, from Gomez Farias; 173, S^, lukensi; 174, S^. bartolo; 175, S^. reddel Ii;

176, S^. moisii; 177, S^. davisi; 178, S^. mulaiki; 179, S^, pallidus;

180, S^. cookei; 181, S^, portoricensis from Cueva Cerro Hueco, Chiapas;

182, S, mitchelli. Scale: 1 cm = ,06 mm. 171 Figs, 183-190.--Female's spermathecae of the mexicanus group:

183, 184, S^, cookei; 185-187, S^. mexicanus from various localities:

185, Nacimiento del Rio Frio; 186, Cueva Chica; 187, the type locality;

188, S^. mitchelli; 189, 190, S_, reddel Ii: 189, from Cueva de los

Vampiros; 190, from the type locality. Scale: 1 cm = ,04 mm.

Figs. 191-198,--Female's spermathecae of the mexicanus group:

191, 192, ^. lukensi: 191, from the type locality; 192, from Cueva de Cuarteles; 193, S., bartolo; 194, OTU #2; 195, OTU #11; 196, S. pallidus; 197, S^. moisii; 198, OTU #1. Scale: 1 cm = .04 mm.

Figs. 199-211.--Parts of schizomids of the mexicanus group:

109-206, female's spermathecae of S_. portoricensis from various localities: 199, the type locality; 200, Cave Bellamar, Cuba;

201, St, Catherine Parish, Jamaica; 202, Dade County, Florida;

203, Guayaquil, Ecuador; 204, Santa Cruz Island, Galapagos Islands;

205, Actun Xkyc, Yucatan; 206, Cueva Cerro Hueco, Chiapas; 207-211, male's pedipalps: 207-210, S^, mexicanus from various localities:

207, right, lateral view from the type locality; 208, dorsal view of tibia and tarsus-basitarsus from the type locality; 209, right, lateral view from Gomez Farias; 210, right, lateral view from

Sotano del Tigre; 211, right, lateral view of S^, davisi. Scale:

Figs, 199-206, 1 cm = ,04 mm; Fig, 207, 1 cm = ,12 mm; Figs, 208-

211, 1 cm = .06 mm. 225 Figs. 212-215.—Male's pedipalps of S_, mitchelli: 212, right, lateral view; 213, right, mesal view of tibia and tarsus-basitarsus only; 214, right, lateral view; 215, left, dorsal view. Scale: 1 cm

= ,06 mm. 227 Table 11.--Measurements of the members of the mexicanus group:

1, one female, OTU #1; 2, one female, OTU #2; 3, one male, S^, mulaiki;

4, one male, S^, bartolo; 5, one female, S^, bartolo; 6, one male, S^. lukensi; 7, one female, ^, lukensi; 8, one male, S^, davisi; 9, one male, S^, reddel Ii; 10, one female, S^, reddel Ii; 11, nine males, S^. mexicanus; 12, 12 females, S^. mexicanus; 13, two males, S^. pallidas;

14, one female, S^, pallidus; 15, one female, OTU #11; 16, one male,

S^, moisii; 17, one female, S_, moisii; 18, two males, S^. cookei; 19, three females, S^. cookei; 20, eight males, S^, mitchelli; 21, 12 females,

5, mitchelli. 229

TABLE 11

Taxa n 1 2 3 4 5 6 7 8 9

1 1,14 1,14 ,98 ,99 1.04 1.14 1,34 1,18 1,12 2 .37 ,25 ,26 ,33 ,28 .45 ,35 ,35 ,49

3 - - ,22 ,21 - .18 - ,23 ,20

4 1.46 ,86 ,90 1,16 1,09 1,50 1.30 1,03 1,13

5 1.70 ,97 1,08 1,47 1,41 1,94 1.16 1.29 1,57

6 1,28 ,73 .83 1,13 1,11 1,49 1,25 .97 1.13

7 1,07 ,72 .74 ,95 ,86 ,97 1,00 ,83 1.02 8 .97 ,65 ,62 ,70 ,68 ,90 ,90 ,73 .82 9 .49 .39 .36 ,38 ,32 ,32 ,41 .45 .40 10 ,50 .40 .40 ,50 ,53 ,57 ,62 ,46 .59 11 ,52 .37 ,35 .43 .37 ,51 ,45 ,41 .48 12 ,86 .61 ,56 .63 .60 ,79 ,78 ,64 .70 13 ,40 ,25 .25 .27 .28 ,35 ,34 ,33 .29 14 .57 ,32 .33 ,41 .38 ,53 ,53 ,36 .44 15 ,61 .39 ,37 .47 ,38 ,54 ,53 ,43 .48

16 1.31 .97 ,84 1.05 1,00 1,22 1,19 ,97 1.05 17 .47 .44 .35 .35 ,36 ,47 ,42 ,46 .38 18 1.00 ,63 .61 .78 .77 ,90 ,81 ,72 ,81 19 .93 .58 ,52 ,68 .68 ,79 ,74 ,60 ,75 230

TABLE 11 (continued)

Taxa

n 10 11 12 13 14 15 16

1 1,13 1.14-1.28 1.20-1.31 1.21-1,37 1,44 .99 1.04 2 ,30 ,39-,40 .31-.35 ,49-,53 ,45 ,24 .37

3 - ,20-,22 - ,25-,26 - - ,26

4 1.02 1,23-1.39 1.10-1.20 1,64-1,86 1,73 ,85 1,32

5 1.26 1.43-1.71 1.33-1.42 2,11-2,12 2,08 ,97 1,78

6 .90 1.05-1,61 .94-1.02 1,52-1,53 1.60 .71 1,30

7 .81 ,92-,97 .84-.97 1,20-1,22 1,11 .67 ,95

8 ,73 ,90-.97 .82-,86 1,07-1.09 1,15 .59 ,78

9 ,41 .49-.54 .45-.48 .52-.59 ,63 .34 ,42

10 ,47 ,62-.67 .55-.59 .80-,80 ,80 .35 ,52

11 .39 ,48-,52 .43-.49 .55-.58 ,62 ,35 .50

12 .64 .80-.82 .74-.89 .97-.99 1,06 ,54 .62

13 .30 ,31-,40 .35-,49 .46-.61 ,45 ,23 .28

14 .37 ,47-,50 ,42-,59 .59-.78 ,62 .26 .35

15 ,44 .54-.58 .48-.51 .63-.65 .68 .35 .40

16 1,02 1.22-1.30 1.15-1.21 1.50-1.53 .81 .87 1.18

17 .40 .52-.59 .50-.54 .60-.64 .62 .41 .51

18 .71 .89-,92 .78-.87 1.09-1,11 1.21 .57 .82

19 .62 ,77-,82 .70-.76 .93-,95 1.00 .54 .67 231

TABLE 11 (continued)

Taxa n 17 18 19 20 21

1 1.13 1,12-1.16 1,17-1,23 1.04-1.16 1.04-1.16

2 .26 .43-.45 ,37-,38 .40-.45 .28-.31

3 - ,45-,45 - ,35-,39 -

4 1,11 1,14-1,32 1,09-1,18 1,10-1,24 .98-1.05

5 1,42 1,37-1.55 1,34-1,39 1.34-1.49 1.12-1.24

6 1,00 1.03-1,15 1,02-1,07 1.02-1.09 .87-.92

7 ,86 .92-,97 ,81-,89 .87-.95 .76-.81

8 .70 .84-,96 ,79-.90 ,74-,82 .69-,76

9 .37 .45-,51 .44-.50 ,42-,49 ,39-,44

10 .46 ,59-,66 .49-.59 ,51-,58 .41-,49

11 ,44 ,50-,52 .45-.54 ,48-.48 .34-.43

12 .67 ,74-,80 .76-.78 .68-.74 .61-.68

13 .31 .30-.39 .35-.39 .30-.34 .29-.32

14 .41 .41-,49 .44-.48 .38-.45 .31-,38

15 .42 ,51-,57 .51-.55 .45-,52 .42-.45

16 1.06 1.09-1,24 1.13-1.13 1,04-1.17 .97-1.05

17 .43 .41-,49 .46-.51 .45-.50 .41-.47

18 ,80 ,82-,90 .84-.89 .74-,83 ,69-.77

19 ,65 ,71-,83 .70-.76 "- ,65-. 72 .59-.64 232

The pecki group

Members of this group are characterized by large size

(1.33-1.76, carapacal length). The color is brownish. The eyespots vary from absent to indistinct to distinct. Suspected troglobites

lack eyespots. The carapace has two or three pairs of dorsal and

two apical setae, the median pair of the dorsal setae of those

species with three pairs are invariably the smallest. The abdc~en

is never attenuated. Males, as far as is known, lack the

posterodorsal process. The males' flagella are large and bulbous.

The females' flagella are always long (,37-.47) and are composed of

three articles, and may be terminally expanded. The females'

spermathecae are characterized by the median pair usually being much

longer than the lateral pair. The laterals are sometimes absent,

and sometimes nearly as long as the medians,, They are withou~ ~uch

terminal enlargement or localized sclerotization. The pedipalps are

very stout in both sexes and do not exhibit dimorphism. The

pedipalpal claws are large, usually 2/3 and up to 3/4 the length of

the tarsus-basitarsus.

Distribution.--Mexico: Veracruz, Oaxaca, Tabasco, Chiapas.

Belize, Guatemala.

Subordinate taxa,--OTU #8; firstmani complex: i, firstniani,

OTU #2, S^. sp, (cf,) sbordonii; pecki complex: S^. pecki, S^,

ayatemaXensis, OTU ^6, OTU #7. Table 12.—Comparisons of the members of the pecki group. For explanation of the characters see Phylogeny. 234

guatem­ first­ sbor­ pecki OTU #6 OTU #7 OTU #8 CHARACTER mani OTU #2 donii alensis

DORSAL 3 3 3 2 2 2 2 3 SETAE

STERNAL 8 9 9 8 10 13 9 9 SETAE

Indis­ Indis­ indis­ Indis­ distinct absent distinct EYESPOTS absent tinct tinct tinct tinct

SPERMA­ laterals M 2X L M 3X L M 2X L M 4X L H += L M ±= L M 2X L THECAE absent

CARAPACAL 1.40 1.54 1.52 1.58 1.33 1.41 1.76 1.49 LENGTH

DISTAL EXP. strong strong slight slight 7 slight none none FEM. FLAG.

LENGTH FEM. .38 .47 .39 .46 7 .41 .47 .45 FLAGELLUM

PIT MALE absent 7 ? single 7 7 7 7 FLAGELLUM 235

Schizomus firstmani Rowland, 1973

(Figs. 217, 219, 221, 226, 227; Tables 12, 13; Map 4)

Schizomus firstmani Rowland, 1973a:7, 15, 16-19; Rowland, 1973c:136.

Holotype,--An adult male, taken in Grutas de Atoyac, 2 km E

Atoyac, Veracruz, Mexico, on 24 December 1972, by D. McKenzie (AMNH,

examined).

Allotype,--An adult female, taken in Grutas de Atoyac, on

6 August 1969, by S, and J, Peck (AMNH, examined),

Paratypes,--Two adult females and three immatures taken with

the allotype (AMNH, examined); an immature, taken with the holotype

(AMNH, examined); an adult female and three immatures, taken in Grutas

de Atoyac, on 22 August 1965, by J, Reddell, J, Fish, W, Bell (AMNH,

examined).

Description.--Male. Color brownish. Carapace with three pairs

of dorsal, medians very small, and two apical setae, Eyespots absent. Anterior sternum with eight bifid setae. Abdominal terga I-VII with

two setae, terga VIII-IX with four setae, segment XII with no evidence

of posterodorsal process. Vestigial stigmata darker than sterna,

Flagellum compressed laterally, expanded distally, with complex

sculpturing. Pedipalpal trochanter produced slightly distally,

tarsal-basitarsal spurs about 1/3, claw about 2/3 length of tarsus-

basitarsus, Tarsal-basitarsal segments of leg I of the following

approximate proportions: 53-8-10-10-10-13-24, Other leg segment

measurements given in Table 13, 236

Female, Flagellum composed of three articles, expanded greatly distally. Median spermathecae only present, long slender, slightly divergent apically, terminating in slight bulbs.

Comparisons,--The female of S^, firstmani is very similar to that of OTU #2, They are perhaps distinguishable by the form of the female's spermathecae, in that OTU #2 has lateral spermathecae. In addition OTU #2 has a much longer flagellum (,47) compared to S^, firstmani (.38), OTU #2 and S^, firstmani are distinct from other members of the group, and other American schizomids, in having the female's flagellum distally greatly enlarged. On this basis the two taxa are considered closely related.

Distribution,--This species is known from caves in Veracruz and Oaxaca, Mexico,

Remarks,--While this species is apparently troglobitic I have here included under its name another population of very similar organisms from Cueva del Nacimiento del Rio San Antonio, Oaxaca, a cave of about 30 km distance from the type locality. They are morphologically distinct in no major way, but it is doubtful that genetic interchange is possible between them. They are probably recently isolated sister species of a once wider ranging population.

S_, firstmani shares Grutas de Atoyac with an epigean member of the pecki group which I have tentatively placed under the name S^, sbordonii. 237

Additional records,--Oaxaca: Cueva del Nacimiento del Rio San

Antonio, 10 km S Acatlan, 26 December 1972, J, Reddell, D, McKenzie,

M, McKenzie, S, Murphy, one female, three immatures (TTU),

Schizomus sp,, OTU #2

(Figs, 222, 223, 229, 234; Tables 12, 13; Map 4)

Specimens examined,--Three adult females, six immatures, taken in Grutas de Zapaluta, 4 mi SE Zapaluta, Chiapas, Mexico, on

28 August 1972, by R, Mitchell, J, Cooke (TTU); an adult female, taken in Grutas de Zapaluta, on 19 July 1950, by C, and M, Goodnight

(AMNH); an adult female, taken in Grutas de Zapaluta, on 20 August

1967, by J, Reddell, J, Fish, T, Evans (TTU); an adult female, one immature, taken in Sumidero del Camino, 10 mi NE Comitan, Chiapas, on 22 August 1967, by J, Reddell, J, Fish (TTU); an immature, taken in Cueva Chica de Hun Chabin, near Comitan, Chiapas, on 21 August

1967, by J, Reddell (TTU),

Description,--Male unknown. Female, Color brownish.

Carapace with three pairs of dorsal and two apical setae. Eyespots indistinct. Anterior sternum with nine bifid setae. Abdominal terga

I-VII with two setae, terga VIII-IX with four setae. Vestigial stigmata darker than sterna. Flagellum composed of three articles, markedly expanded distally. Pedipalpal trochanter greatly produced distally, tarsal-basitarsal spurs about 1/4, claw about 2/3 length of tarsus-basitarsus. Tarsal-basitarsal segments of leg I of the following approximate proportions: 51-8-10-8-11-10-21. Other leg 238 segment measurements given in Table 13, Median spermathecae twice length of laterals, both divergent, medians curved, with slight bulbs, but no special sclerotization.

Comparisons,--See under S^, firstmani,

Distribution,--This taxon is known from various caves near

Comitan and Zapaluta, Chiapas, Mexico,

Remarks,--OTU #2 is known only from caves, but does not show the advanced troglobite facies of the other species of the pecki group. It will probably be found to occur in epigean habitats near the caves in which it is found.

Schizomus sp,, cf, sbordonii Brignoli, 1973

(Fig, 225; Tables 12, 13; Map 4)

Schizomus sbordonii Brignoli, 1973:7, 8, 9; Rowland, 1973c:135, 136;

Brignoli, 1974:143, 146, 147-149, 151,

Specimen examined.--An adult female, taken in Cueva de Atoyac,

Atoyac, Veracruz, Mexico, on 6 August 1969, by S. and J. Peck (TTU).

Holotype.--An adult female, taken in Cueva de Ojo de Agua

Grande, near Cordoba, Veracruz, on 5 November 1969, by V, Sbordonii

(IZR, not examined),

Description of female from Cueva de Atoyac,--Color brownish.

Carapace with three pairs of dorsal and two apical setae. Eyespots distinctly round. Anterior sternum with nine bifid setae. Abdominal terga I-V, VII with two setae, terga VI with three setae, terga

VIII-IX with four setae. Vestigial stigmata darker than sterna. 239

Flagellum long, composed of three articles, slightly expanded distally, Pedipalpal trochanter produced distally, tarsal-basitarsal spurs about 1/4, claw about 1/2 length of tarsus-basitarsus,

Tarsal-basitarsal segments of leg I of the following approximate proportions: 48-8-9-10-10-11-23, Other leg segment measurements given in Table 13, Median spermathecae three times longer than laterals, both divergent, medians with apical half sclerotized, medians and laterals without terminal bulbs.

Comparisons.--The specimen here studied seems to be very similar to S^, firstmani and OTU #2 in most respects, but differs from them in having very distinct and round eyespots, only a slight expansion of the flagellum, and is more darkly pigmented.

Distribution,--If the specimen studied is truly representative of S^, sbordonii, then this species is known from Cueva de Atoyac,

Atoyac, and Cueva de Ojo de Agua Grande, near Cordoba, Veracrjz,

Mexico,

Remarks,--I have included the collection from Cueva de Atoyac under the name S^, sbordonii for two reasons. The description and illustrations given by Brignoli (1973, 1974), while really inadequate to recognize any species, do not contraindicate the specimen I studied. More importantly, the very close geographic proximity and most probable epigean nature of these populations suggest that they are conspecific.

The types of S^, firstmani were collected with the specimen I studied, but could have come from a remote part of the cave. The 240 latter is definitely a darkly pigmented, epigean representative, while

S. firstmani is an unmistakable troglobite,

Schizomus pecki Rowland, 1973

(Figs, 216, 218, 231; Tables 12, 13; Map 4)

Schizomus pecki Rowland, 1973a:7, 16, 19-23; Rowland, 1973c:136,

Holotype,--An adult male, taken in Las Grutas del Cocona,

2 mi NE Teapa, Tabasco, Mexico, on 1 August 1948, by C, Goodnight

(AMNH, examined).

Allotype,--An adult female, taken in Las Grutas del Cocona, on 29 November 1971, by D, McKenzie (AMNH, examined),

Paratype,--An adult female, taken with the allotype.

Description,--Male, Color brownish. Carapace with two pairs of dorsal and two apical setae, Eyespots indistinct. Anterior sternum with eight entire setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segment XII without evidence of posterodorsal process. Vestigial stigmata darker than sterna.

Flagellum lanceolate, with a median depression following a single median gentle elevation, Pedipalpal trochanter produced distally,

tarsal-basitarsal spurs about 1/5, claw about 2/3 length of tarsus-

basitarsus, Tarsal-basitarsal segments of leg I of the following

approximate proportions: 62-8-10-9-9-19, Other leg segment measurements given in Table 13.

Female. Flagellum composed of three articles, slightly

expanded distally. Median spermathecae twice length of laterals. 241 both divergent, medians curved, but without terminal bulbs, both pair with light sclerotization along distal half of both lobes.

Comparisons.--S^. pecki is similar to three other middle

American members of the pecki group. The two pairs of dorsal carapacal setae and the long (,40), slightly distally expanded flagellum of the female readily distinguish this species from the others, S^, pecki is characterized by having a distinctly smaller pedipalpal claw than females of OTU #7, and a distinctly larger claw than OTU #6 and S^, guatemalensis females. The forms of the spermathecae are diverse .in this assemblage, S^, pecki and S^, guatemalensis are similar in that the medians are considerably larger than the laterals, more so in S^. guatemalensis, and they are nearly equal in OTU #6 and OTU #7,

Distribution,--This species is known only from Las Grutas del

Cocona, near Teapa, Tabasco, Mexico.

Remarks.--S^. pecki is apparently a troglobite, but shares occupancy of Las Grutas del Cocona with S^. trilobatus, a brasiliensis group trogloxene. For a further discussion of their relationships see under the latter species.

Additional records,—Las Grutas del Cocona, 24 July 1973, J,

Reddell, J, Rowland, three females, three immatures (TTU).

Schizomus guatemalensis Chamberlin, 1922

(Fig. 228; Tables 12, 13; Map 4)

Schizomus guatemalensis Chamberlin, 1922:12; Mello-Leitao, 1931:17;

Giltay, 1935:6; Gertsch, 1941:14; Takashima, 1943:93. 242

Holotype.--An adult female, taken at San Rafael, Guatemala

(MCZ, examined).

Description.--Male unknown. Female. Color brownish.

Carapace with two pairs of dorsal and two apical setae. Eyespots indistinct. Anterior sternum with 10 bifid setae. Abdominal terga

I-VII with two setae, terga VIII-IX with four setae. Vestigial stigmata darker than sterna. Flagellum missing. Pedipalpal trochanter produced distally, tarsal-basitarsal spurs about 1/5, claw about 1/2 length of tarsus-basitarsus. Tarsal-basitarsal segments of leg I of the following approximate proportions: 40-6-8-7-9-9-20. Other leg segment measurements given in Table 13. Median spermathecae long and slender, strongly curved outwardly, lightly sclerotized along distal 2/3, terminating in a slight bulb, laterals about 1/4 as long, straight, but directed diagonally outward, no terminal bulb or special sclerotization.

Comparisons.--See under S^. pecki.

Distribution,--This species is known only from San Rafael,

Guatemala, Also see Remarks,

Remarks,--The locality assigned by Chamberlin cannot be associated with certainty to any of the several communities of the name San Rafael in Guatemala, S^, guatemalensis shows none of the troglobite facies common to other members of the group and is apparently representative of an epigean population. 243

Schizomus sp., OTU #6

(Fig, 232; Tables 12, 13; Map 4)

Specimen examined,--An adult female, taken in St. Herman's

Cave, 400 ft,. Caves Branch, British Honduras, between 23 July and

21 August 1972, by S. and J, Peck (AMNH).

Description.--Male unknown. Female. Color brownish.

Carapace with two pairs of dorsal and two apical setae. Eyespots absent. Anterior sternum with 13 bifid setae. Abdominal terga

I-VII with two setae, terga VIII-IX with four setae. Vestigial stigmata darker than sterna, Flagellum composed of three articles, long, slightly distally expanded, Pedipalpal trochanter produced distally, tarsal-basitarsal spurs about 1/4, claw about 1/2 length of tarsus-basitarsus. Tarsal-basitarsal segments of leg I of the following approximate proportions: 45-8-9-10-11-20. Other leg segment measurements given in Table 13. Median and lateral spermathecae about same size, slightly convergent, gradually expanded distally; no special sclerotization or apical bulbs.

Comparisons.--See under S^. pecki.

Distribution.--This taxon is known only from St. Herman's

Cave, Caves Branch, British Honduras.

Remarks,--The very light pigmentation and lack of eyespots in this species suggests that it may be an obligate cavernicole, as may be S^, firstmani, OTU #8, and S^, pecki of this group. 244

Schizomus sp., OTU #7

(Figs. 224, 230; Tables 12, 13; Map 4)

Specimens examined.--An adult female, taken at Las Ruinas de

Palenque, near Palenque, Chiapas, Mexico, during July 1948 by C. and

M, Goodnight (AMNH); two immatures, taken at Las Ruinas de Palenque, on 25 July 1973, by J, Reddell (TTU),

Description.--Male unknown. Female. Color brownish.

Carapace with two pairs of dorsal and two apical setae. Eyespots

indistinct. Anterior sternum with nine bifid setae. Abdominal terga

I-VII with two setae, terga VIII-IX with four setae. Vestigial

stigmata darker than sterna, Flagellum broken, but appearing not to

be greatly expanded distally. Pedipalpal trochanter produced distally,

tarsal-basitarsal spurs about 1/3, claw about 3/4 length of tarsus-

basitarsus. Tarsal-basitarsal segments of leg I of the following

approximate proportions: 50-7-10-9-9-11-24. Other leg segment measurements given in Table 13. Median spermathecae slightly smaller

than laterals, both convergent, not sclerotized, not terminating in

bulbs.

Comparisons.--S^. pecki seems most closely related to this

taxon in that they share a similarity in development of the

spermathecae, two pairs of dorsal carapacal setae, development of

eyespots, but the pedipalpal claw is relatively shorter in S^. pecki,

and S^. pecki is smaller in carapacal length. Also see comparisons

under S^. pecki. 245

Distribution,--This taxon is known only from Las Ruinas de

Palenque, near Palenque, Chiapas, Mexico.

Remarks.--J, Reddell reported (personal communication) that he found these under rocks on the deeply shaded canyon walls above the ruins.

Schizomus sp,, OTU #8

(Fig, 233; Tables 12, 13; Map 4)

Specimens examined,--An adult female and one immature, taken in Grutas de Monteflor, 6 km N Valle Nacional, Oaxaca, Mexico, on 28

December 1972, by J, Reddell, D. McKenzie, M, McKenzie, and S, Murphy

(TTU).

Description,--Male unknown. Female, Color brownish.

Carapace with three pairs of dorsal, the medians very small, and two apical setae. Eyespots distinct. Anterior sternum with nine bifid setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae. Vestigial stigmata darker than sterna. Flagellum composed of four articles, not distally expanded, elongate, Pedipalpal trochanter produced slightly distally, tarsal-basitarsal spurs about 1/4, claw about 2/3 length of tarsus-basitarsus, Tarsal-basitarsal segments of leg I of the following approximate proportions: 41-7-8-9-10-10-18,

Other leg segment measurements given in Table 13, Median and lateral spermathecae slightly convergent, medians about twice as long as laterals, both slightly more sclerotized apically than basally, medians terminating in vague bulbs. Figs, 216-221,—Parts of schizomids of the pecki group: 216-

219, male's flagella: 216, 217, dorsal views: 216, S_. pecki; 217,

S^. firstmani; 218, 219, lateral views: 218, S^. pecki; 219, S^. firstmani; 220, 221, lateral views of right pedipalps: 220, S^. firstmani; 221, S^, pecki. Scale: Figs. 216-219, 1 cm = .06 mm;

Figs. 220, 221, 1 cm = ,25 mm.

Figs, 222-228,--Female's spermathecae of the pecki group: 222,

223, OTU #2: 222, from the type locality; 223, from near Comitan;

224, OTU #7; 225, S_. (cf.) sbordoni i; 226, 227, S^, firstmani: 226, from the type locality; 227, from Cueva del Nacimiento del Rio San

Antonio; 228, S^, guatemalensis. Scale: 1 cm = .04 mm.

Figs, 229-234,--Parts of schizomids of the pecki group: 229,

230, lateral views of female's left pedipalps: 229, OTU #2; 230,

OTU #7; 231-233, speniiathecae: 231, S_, pecki; 232, OTU #6, 233,

OTU #8; 234, lateral view of flagellum of OTU #2, Scale: Figs, 229,

230, 1 cm = .12 mm; Figs. 231-233, 1 cm = .04 mm; Fig, 234, 1 cm =

,06 mm. 251 Table 13.--Measurements of the members of the pecki group:

1, one male, S^, firstmani; 2, one female, S_, firstmani; 3, three females, OTU #2; 4, one female, S^, (cf,) sbordonii; 5, one male,

S^. pecki; 6, two females, S^. pecki; 7, one female, S^. guatemalensis;

8, one female, OTU #6; 9, one female, OTU #7; 10, one female,

OTU T8. 253

TABLE 13

Taxa n 1 2 3 4 5 6 7

1 1.25 1,40 1.46-1.54 1,52 1.31 1,55-1,58 1,33

2 .46 ,38 ,46-,47 ,37 .60 ,46-,47 -

3 .29 - • - ,29 - -

4 1.85 1,78 1,81-1,87 1.67 2,24 2,05-2,44 1.32

5 2.12 2.13 2,08-2,18 2.20 3,74 2,50-2,51 1,56

6 1.48 1.45 1.46-1.50 1.69 2,12 1,88-1,94 1,12

7 1.20 1.20 1,20-1.24 1.19 1,25 1,15-1,16 ,99

8 1,15 1.18 1.28-1.35 1.22 1.21 1,20-1,36 ,98

9 ,56 ,62 .65-.73 .68 ,57 ,61-,70 ,51

10 ,80 ,80 .86-.94 ,84 ,90 ,85-,85 .61

11 ,59 ,63 .72-.76 .67 ,79 ,73-,77 .54

12 1.00 1,07 1.20-1.23 1.07 1.10 1,17-1,20 .87

13 .40 ,52 .56-.64 .55 ,50 .55-,59 .43

14 .69 ,67 .80-.84 .72 ,75 .71-.74 .50

15 ,63 ,73 .78-.86 .73 ,80 ,83-,84 .61

16 1.63 1.65 1.78-1.86 1.04 1,96 1,82-1,93 1.34

17 .62 .68 .76-.79 .75 ,65 ,63-,83 .62

18 1.18 1,12 1.28-1.31 1.20 1,36 1,30-1,37 ,92

19 1.02 1.04 1,15-1,19 1.01 1,14 1,04-1,11 ,85 254

TABLE 13 (continued)

Taxa

n I 8 9 10

1 •1-41 1,76 1,49

2 •41 ,47 .45 3

4 1-53 2.04 1.57

5 1.90 2.50 1.90

6 1.44 1.85 1,41

7 1-13 1,20 1,03

8 1-06 1.45 1.09

9 .54 ,77 ,62

10 .71 1,00 .72

11 •55 .75 .57

12 .93 1.22 .95

13 .44 ,58 .45

14 .55 ,75 ,54

15 .63 .82 .62

16 1.48 1.90 1.49

17 .62 .80 .67

18 1.02 1.38 1.08

19 .89 1.16 .91 255

Comparisons,--This taxon is unique among members of the pecki group in that the female's flagellum is not at all expanded distally.

It shares with several other species in this group possession of three pairs of dorsal carapacal setae, but is the only one among them with sclerotized spermathecal tips.

Distribution,--This taxon is known only from Grutas de

Monteflor, near Valle Nacional, Oaxaca, Mexico.

Remarks.--This taxon, like many of the pecki group, shares its cave environment with another schizomid. In this case Grutas de

Monteflor is cohabited by OTU #8 and S^. moisii, a mexicanus group member of dark pigmentation and undoubted facultative cave occupancy.

The distinct eyespots and moderate pigmentation of OTU #8 leads me to believe that it is also a facultative cavernicole.

The goodnightorum group

Members of this group are characterized by small to large size (,89-1,42 carapacal length). The color is brownish. The eyespots are distinct or indistinct, and are with irregular or round margins. The carapace has three or four pairs of dorsal and two apical setae. Abdominal segments VII or VIII to XII may be slightly to extremely attenuated in the males. No posterodorsal abdominal process is evident. The male's flagellum is always elongate, narrow, and apically slender, and has either double or a single median pit.

The female's flagellum is moderate in length (.22-.31) and is composed of three articles. The female's spermathecae are characterized by the Table 14,--Comparisons of the members of the goodnightorum group. For explanation of the characters see Phylogeny, 257

CHARACTER goodni- orth- lance- silvino ghtorum opiax olatus

DORSAL SETAE

STERNAL SETAE 10 10 11 10

EYESPOTS ^tlnct "iistlnct distinct distinct

SPERMA­ THECAE M 5X L M 2X L

CARAPACE .89 1.04 1.42 1.05 LENGTH

LENGTH FEM. FUGELLUM .22 .31

ABDOMINAL ELONGATION 7-12 8-12 7-12 8-12

PIT MALE single double double double FLAGELLUM 258 median pair being much longer than the laterals, with small or no terminal bulbs, and with slight sclerotization. The pedipalps are not sexually dimorphic.

Distribution,--Mexico: Veracruz, Chiapas, Yucatan; Guatemala,

Subordinate taxa,--S^, goodnightorum, S^. orthoplax, S^, lanceolatus, S^, silvino,

Schizomus goodnightorum (Rowland), 1973 NEW COMBINATION

(Figs, 238, 239, 241, 242, 247; Tables 14, 15; Map 4)

Heteroschizomus goodnightorum Rowland, 1973a:2-6; Rowland, 1973b:202,

Holotype,--An adult male, taken at Chichen Itza, Yucatan,

Mexico, in June 1948 by C, Goodnight (AMNH, examined),

Paratype,--An adult male, taken with the holotype (AMNH, examined).

Female.--An adult, taken at Chichen Itza, Yucatan, Mexico, in June 1948 by C. and M. Goodnight (AMNH, examined).

Description.--Male, Color brownish. Carapace with three pairs of dorsal and two apical setae, Eyespots indistinct. Anterior sternum with 10 bifid setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segments VII-XII extremely elongate, segment XII with no evidence of posterodorsal process. Vestigial stigmata lighter than sterna, Flagellum long and extremely narrow, with a median pit, Pedipalpal trochanter produced distally, tarsal-basitarsal spurs about 1/5, claw about 1/3 length of tarsus-basitarsus. Leg segment measurements given in Table 15, 259

Female, Flagellum composed of three articles, Spermathecae with medians several times longer than laterals, both divergent, the

laterals thicker than medians, the latter terminating in slightly

sclerotized, vaguely defined bulbs.

Comparisons,--S^, goodnightorum has a much greater elongation

of distal abdominal segments in the males, and one fewer pair of

dorsal carapacal setae than other members of this group. Males of

the other species have a pair of dorsal flagellar depressions, while

S_. goodnightorum has a single depression. The male of this species is further distinguishable from other males with any degree of

attenuation of distal abdominal segments outside the goodnightorum

group by its lack of a posterodorsal abdominal process. The median

spermathecae of ^, goodnightorum are much longer than those of S^,

silvino, the only other species in this group known by females.

Distribution,--This species is known only from Chichen Itza,

Yucatan, Mexico,

Remarks,--The placement of this species in the genus Schizomus

was, in part, necessitated by the discovery of several other species

with a considerably less, but none the less distinct, attenuation of

distal abdominal segments. Since the original description a female,

undoubtedly referable to this species, has also become available for

study. It has thus become apparent that while a unique development

of the abdomen obtains in the males of this species, the females are

in my opinion without peculiarities such that would distinguish them

significantly from the females now contained in the genus Schizomus, 260

Variation,--The male paratype is smaller than the holotype and exhibits a lesser degree of abdominal attenuation. As was noted in the original description the paratype contained a nematode parasite in the abdominal cavity which may account for its smaller size, but variation of this magnitude is not unusual in other species,

Schizomus orthoplax Rowland, 1973

(Figs, 236, 240, 244; Tables 14, 15; Map 4) Schizomus orthoplax Rowland, 1973a:6, 10-13; Rowland, 1973c:135.

Holotype,--An adult male, taken at the Finca Cuauhtemoc, Chiapas, Mexico, on 8 May 1950, by C, and M, Goodnight (AMNH, examined). Description.--Male, Color brownish. Carapace with four pairs of dorsal and two apical setae, Eyespots distinctly round. Anterior sternum with 10 bifid setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segment XII with no evidence of posterodorsal process. Vestigial stigmata darker than sterna, Flagellum lanceolate, with a pair of median pits, on otherwise flat dorsal surface, Pedipalpal trochanter produced distally slightly, tarsal-basitarsal spurs about 1/4, claw about 1/2 length of tarsus-basitarsus, Tarsal-basitarsal segments of leg I of the following approximate proportions: 35-4-7-6-7-7-16, Other leg segment measurements given in Table 15,

Female unknown. 261

Comparisons,--This species is most similar to S^, silvino and

S. lanceolatus. They are best distinguished on the basis of the males' flagella, though the carapacal lengths also differ among these species.

The flagellum is by far the longest in S^, lanceolatus and by far the shortest in S^, orthoplax and is apparently intermediate in S^, silvino.

See also under S^, goodnightorum.

Distribution,--This species is known only from the Finca

Cuauhtemoc, Chiapas, Mexico.

Additional records.--Chiapas: Finca Cuauhtemoc, 8 June 1950,

C. and M. Goodnight, two immatures (AMNH),

Schizomus lanceolatus Rowland, 1975

(Figs, 235, 243; Tables 14, 15; Map 4)

Schizomus lanceolatus Rowland, 1975:7, 15-16, 17,

Holotype,--An adult male, taken in Cueva del Diablo, near

Ciudad Mendoza, Veracruz, Mexico, on 7 March 1973, by J, Reddell (TTU, examined).

Description,--Male, Color brownish. Carapace with four pairs of dorsal and two apical setae. Eyespots distinct, irregular.

Anterior sternum with 11 bifid setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae, abdominal segments VII-XII attenuate, segment XII with no evidence of posterodorsal process.

Vestigial stigmata slightly darker than sterna. Flagellum lanceolate, with pair of median pits on otherwise flat dorsal surface, Pedipalpal trochanter produced distally, tarsal-basitarsal spurs about 1/4, claw 262 about 1/2 length of tarsus-basitarsus, Tarsal-basitarsal segments of leg I of the following approximate proportions: 57-9-9-9-10-9-21,

Other leg segment measurements given in Table 15,

Female unknown.

Comparisons.--See under S^, orthoplax and S^, goodnightorum.

Distribution.--This species is known only from Cueva del

Diablo, near Ciudad Mendoza, Veracruz, Mexico.

Remarks.--While this species was collected from a cave, it is sufficiently darkly pigmented that I suspect it is a facultative cavernicole, probably also occuring in moist areas outside the cave.

Schizomus silvino, new species

(Figs. 237, 245, 246; Tables 14, 15; Map 4)

Holotype.--An adult male, taken in Gruta de Silvino, 34 km

W Puerto Barios, Izabal, Guatemala, between 20-22 August 1969, by

S. and J. Peck (AMNH)

Allotype.--An adult female, taken with the holotype (AMNH),

Paratypes,--Three adult males, six adult females, and six

immatures, taken with the holotype (AMNH),

Description,--Male. Color brownish, pale. Carapace with

four pairs of dorsal and two apical setae, Eyespots distinct,

irregular. Anterior sternum with 10 bifid setae. Abdominal terga

I-VII with two setae, terga VIII-IX with four setae, segments VIII-XII

attenuate, segment XII with no evidence of posterodorsal process.

Vestigial stigmata lighter than sterna, Flagellum extremely Figs, 235-240.—Parts of male schizomids of the goodnightorum group: 235-238, dorsal views of flagella: 235, S^. lanceolatus;

236, S^. orthoplax; 237, S_. silvino; 238, S^, goodnightorum; 239, 240, lateral views of right pedipalps: 239, S^. goodnightorum; 240, S^. orthoplax. Scale: Figs, 235-238, 1 cm = ,06 mm; Figs, 239, 240,

1 cm = ,25 mm.

Figs, 241-247.--Parts of schizomids of the goodnightorum group:

241, dorsal view of male S^. goodnightorum, legs and pedipalps past the trochanter omitted; 242-245, lateral views of male's flagella,: 242

S^. goodnightorum; 243, S^. lanceolatus; 244, S^, orthoplax; 245, S_. silvino; 246, 247, female's spermathecae: 246, S^, silvino; 247, S_. goodnightorum. Scale: Fig. 241, 1 cm = .4 mm; Figs, 242, 243, 245,

1 cm = ,12 mm; Fig, 244, 1 cm = ,06 mm; Figs, 246, 247, 1 cm = .04 mm.

^MMMMMMMMMI 266

242 I u

1^

244

245

247

241 Table 15.—Measurements of the members of the goodnightorum group: 1, two males, S^. goodnightorum; 2, one female, S_. goodnightorum; 3, one male, S^. orthoplax; 4, one male, S^. lanceolatus; ; ; ; ;

5, two males, S. silvino; 6, two females, S. silvino. 268

TABLE 15 Taxa

n 1 2 3 4 5 6 1 1.04-1.12 ,89 1,04 1,42 1,11-1.14 1.00-1.05 2 1.04 .22 .57 ,99 ,70-.79 ,29-,31

3 ,22 - ,19 ,27 .20-.20 - 4 1.33-1,67 ,84 1.02 1.80 1,45-1,62 1,05-1,14 5 1.55 ,99 1.24 2,34 1,80-2,01 1,26-1.36 6 1.15 ,72 ,85 1,79 1.03-1.36 ,86-.91

7 - ,60 .82 1.26 .97-.99 .76-.77 8 .83-,93 .58 .60 1,15 .82-,92 .71-.75 9 .40-.52 ,33 .31 ,55 .48-,51 .40-.41 10 .56-,63 ,32 .38 ,82 ,53-,59 ,40-,45 11 ,51-.45 ,30 .38 .67 .49-.53 ,38-,43 12 .77 .51 ,53 1.01 ,71-,79 .61-.65 13 ,30 .22 .20 .50 .33-,36 ,29-,31 14 .41 ,26 ,30 ,57 ,41-,44 ,34-,36 15 .50 ,33 ,37 ,71 ,50-,55 ,40-,47

16 1,16-1,35 ,86 ,94 1,51 1,17-1,31 1,01-1.07

17 ,45-.50 ,40 ,37 .65 .51-.52 ,44-.46

18 .80-,95 .60 ,64 1.07 .77-,86 ,66-.71

19 ,70-,81 ,50 ,51 .92 ,70-,77 ,58-,64 269 elongate, with a pair of median pits on otherwise flat dorsal surface,

Pedipalpal trochanter produced distally, tarsal-basitarsal spurs about

1/5, claw about 2/5 length of tarsus-basitarsus. Tarsal-basitarsal segments of leg I of the following approximate proportions: 44-6-8-7-

8-8-18. Other leg segment measurements given in Table 15.

Female. Flagellum composed of three articles. Median spermathecae two to three times longer than laterals, both very slightly divergent, but neither expanded distally, medians sclerotized slightly on apical half.

Comparisons,--See under S^, orthoplax and S^. goodnightorum.

Distribution,—This species is known only from Gruta de Silvino, near Puerto Barios, Izabal, Guatemala,

Etymology.--The specific name is a noun used in apposition.

Remarks.--While this species is known only from a cave, I am not able to judge whether it is troglobitic. It is paler than most other species, but this is not necessarily a sign of obligate cave existence.

The briggsi group

The members of this group are characterized by moderate to large sized (1,17-1,50, carapacal length) individuals. The color is usually brownish, but may be greenish. The eyespots are indistinct or distinct. The carapace has one or three to four pairs of dorsal and three apical setae. The abdomen is attenuated in males of one species, but the others are normal. The males have a posterodorsal Table 16,—Comparisons of the members of the briggsi group.

For explanation of the characters see Phylogeny. 271

penta­ wessoni borrego­ shoshon­ joshu­ briggsi belkini CHARACTER peltis ensis ensis ensis

DORSAL 3 SETAE 3-4 3 1 3 3 3

STERNAL 14 13 11 13 SETAE 13 13 13

COLOR brown brown brown brown brown green green

HALE short short long long long long PEDIPALP short

SPERMA­ several 7 THECAE several

CARAPACE 1.50 1.30 1.43 1.21 1.41 1.29 1.17 LENGTH

LENGTH FEM. .43 .34 .40 .39 .37 FLAGELLUM .54

ABDOMINAL 7-12 none none none none none none ELONGATION

, ,. tril- penta- trian- hexa- hexa- hexa- SHAPE MALE eiongate ^^^^^^ g^^^^ gy^^^ ggpai gonal gonal FLAGELLUM

PIT MALE absent absent double double single double FLAGELLUM double 272 abdominal process which is always pointed apically. The flagellum is usually globose, but is long and thin in one species, and usually bears some dorsal modifications. The female's flagellum is composed of four articles and is long (.34-.54), The female's spermathecae are characterized by three, four, or several pairs of short, broad lobes which may or may not be sclerotized apically. The pedipalps may be slightly to extremely or not sexually dimorphic. The trochanter is often produced distally, and the femur, patella, and tibia are often elongated. The tibia in males of all species has either a series of heavy spines or a well developed spur apposible to the tarsus- basitarsus.

Distribution.--United States: California, Arizona.

Subordinate taxa.—Pentapeltis complex: S^. pentapeltis; briggsi complex: S^, wessoni, ^, borregoensis, S^, shoshonensis, S^, joshuensis, S^. briggsi, S^. belkini.

Schizomus pentapeltis (Cook), 1899

(Figs, 2, 255, 259, 270; Tables 16, 17; Map 6)

Hubbardia pentapeltis Cook, 1899:253-254, 261,

Trithyreus pentapeltis, Banks, 1900:422; Hansen and Sorensen, 1905:4.

44, 70; Moles, 1917:1-7; Moles, 1921:11; Kishida, 1930:18;

Hilton, 1932:33-34, 45-46; Giltay, 1935:8; Werner, 1935:469;

Gertsch, 1940:1; Takashima, 1943:96; Comstock, 1948:18; Kraus,

1957:245; McDonald and Hogue, 1957:1, 6-7; Briggs and Hom,

1966:270, 273, 274; Hom, 1967:218, 220; Rowland, 1971c:304; 273

Rowland, 1972a:69-74; Rowland, 1972b:5, 8; Rowland, 1972c:153,

155, 156, 159; Rowland, 1973b:195, 195, 202,

Schizomus pentapeltis, Mello Leitao, 1931:18,

Cotypes.--Two adult males, and adult female, taken at Palm

Canyon, near Palm Springs, Riverside County, California, on 13

February and 6 March 1897, by Hubbard (USNM, examined).

Description of a male and female topotype,--Male, Color brownish. Carapace with three pairs of dorsal and three apical setae.

Metapeltidium split, Eyespots distinct, irregular. Anterior sternum with 13 bifid setae. Abdominal terga I-IV with two setae, V-VI with four setae, IX with two setae, VII-VIII with six setae, segment XII with acutely produced, well developed posterodorsal process. Abdominal segments VII-XII attenuated. Vestigial stigmata darker than sterna,

Flagellum long, lanceolate, with a pair of median depressions.

Pedipalpal trochanter produced distally, tarsal-basitarsal spjrs about

1/5, claw about 1/2 length of tarsus-basitarsus, Tarsal-basitarsal segments of leg I of the following approximate proportions:

63-8-12-12-12-11-28, Other leg segment measurements given in Table 17,

Female, Flagellum composed of four articles, Sperrathecae with several small, closely associated lobes, apically scleroiized in localized areas.

Comparisons,—S^. pentapeltis is easily distinguishable from the other species of the group by the elongate flagellum and abdomen of the male. The spermathecal morphology is similar to that of other species, but is characterized by lightly sclerotized apical tubercles. 274

The male's pedipalps are not sexually dimorphic as in other Californian species.

Distribution.--This species is known from the coastal foothills of southernmost California west to the coast and north into the San

Jacinto Mountains, and Los Angeles Basin, An apprently isolated population occurs at an oasis in Palm Canyon on the northeast slope of the San Jacinto Mountains,

Remarks,--This, the first schizomid described from the New

World, has been found to be widespread in California and may occur in Baja California as well. Rowland (1972a) reported the brooding habits and early development of this species.

Variation,--As in other species with elongated abdomen in the males, the abdomen and flagellum seem to vary considerably in the extent to which they are attenuated. Rarely specimens occur which have setational differences from the above description. Four pairs of dorsal carapacal setae may occur, as well as may additional abdominal tergal setae.

Additional records,--Riverside County: Andreas Canyon, 3

March 1956, V, Roth, one male, one female, two immatures (AMNH);

25-27 March 1960, W, Gertsch, W, Ivie, Schrammel, V. Roth, six males,

25 females, 16 immatures (AMNH); 6 April 1966, T. Briggs, K. Hom, four immatures (CAS); 10 January 1971, J, Rowland, one male, five females, five immatures (TTU); 14 January 1971, J, Rowland, two males, six females, three immatures (MCZ); 23 March 1971, J, Rowland, four females, four immatures (LACM); Snow Creek Canyon, 20 March 1954, 275 three females, one immature (AMNH); 13 April 1955, J, Belkin. three females, eight immatures (AMNH); Riverside, 26 March 1960, V. Roth, one male, one female (AMNH); Citrus Experiment Station, Riverside,

21 February 1957, E, Schlinger, one male (AMNH); near Citrus

Experiment Station, Riverside, 1 December 1925, J, Chamberlin, two males, one female (AMNH); Winchester, 22 January 1967, W, Icenogle, one male (LACM); 20 February 1968, W, Icenogle, one male (LACM);

20 February 1969, W, Icenogle, one female (LACM); 18 January 1970,

W. Icenogle, one male (CAS); 17 January 1971, W. Icenogle, one male, one immature (CAS); 19 January 1971, J, Rowland, two males, two females (TTU); 22 January 1971, W, Icenogle, one male (LACM); 24

January 1971, W, Icenogle, one male, three females, one immature

(LACM); 7 February 1971, W, Icenogle, one female (TTU), Orange

County: 11,4 mi SW Lower San Juan Camp, Cleveland National Forest, oak litter, 20 December 1966, A, Jung, 0. Yang, K. Hom, one male, five females, five immatures (CAS); San Clemente, 27 December 1966,

Lum, one immature (CAS). San Diego County: San Diego, March 1970,

B. Kaston,two males, one immature (LACM); El Cajon, 2 March 1969, one male, two females (LACM); 1970, S, Lewis, two males, one female

(LACM); Dripping Springs, near Vail Lake, 6 March 1971, J, Rowland, two males, four females, one immature (AMNH), 276

Schizomus wessoni (Chamberlin), 1939 NEW COMBINATION

(Figs. 254, 258, 263; Tables 16, 17; Map 6)

Trithyreus wessoni Chamberlin, 1939:123-124; Gertsch, 1940:1;

Takashima, 1943:97; Kraus, 1957:245; McDonald and Hogue,

1957:1, 6; Rowland, 1971c:304; Rowland, 1972c:154,

Holotype,--An adult male, taken "under a stone shaded by small bushes and trees growing along the Santa Cruz River" near Tucson,

Arizona, in April 1938, by R, Wesson (AMNH, examined).

Female.--An adult, taken in Tucson, Arizona, by Griswold.

Description.--Male. Color brownish. Carapace with three pairs of dorsal and three apical setae, Eyespots small, indistinct.

Anterior sternum with 14 bifid setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segment XII with acutely produced, well developed posterodorsal process. Vestigial stigmata darker than sterna, Flagellum trilobate, the median lobe arising above the basal two, Pedipalpal trochanter produced distally, thick spines from tibia appose tarsus-basitarsus, Tarsal-basitarsal spurs about 1/6, claw about 1/2 length of tarsus-basitarsus, Tarsal- basitarsal segments of leg I of the following approximate proportions:

37-6-7-7-8-9-21. Other leg segment measurements given in Table 17,

Female, Flagellum composed of four articles, Spermathecae composed of three pairs of small, broad lobes, each with minute terminal elevations, but with no special sclerotization.

Comparisons,—S^, wessoni is distinct from other male members of the group in having a distinctly trilobate flagellum. The 277 pedipalps are very similar in development to those of S^. borregoensis, especially in the development of the mesal tibial spurs which appose the tarsus-basitarsus. The male's flagellum in S^, borregoensis is

distinctly pentagonal and distinct in form from S^. wessoni,

Distribution,—This species is known only from near Tucson,

Arizona.

Remarks,--Long term drying of the Santa Cruz River caused by

agricultural activities upstream from Tucson may have totally

eliminated S^, wessoni from the type locality and perhaps altogether,

A relict population 20 miles south of Ajo, Arizona, may represent an

earlier isolated relative of this species,

Schizomus borregoensis (Briggs and Hom), 1966 NEW COMBINATION

(Figs, 252, 257, 260, 266, 267; Tables 16, 17; Map 6)

Trithyreus borregoensis Briggs and Hom, 1966:270-273; Rowland, i971c:

304, 308-309; Rowland, 1972b:5, 8; Rowland, 1972c:153, 155, 156,

159,

Holotype,--An adult male, taken "under rocks in Palm and

Sycamore debris near stream" in Borrego Palm Canyon, Anza Borrego

State Park, San Diego County, California, on 4 April 1966, by T.

Briggs, K. Horn (CAS, examined).

Allotype.--An adult female, taken with the holotype (CAS,

examined). Paratype.--An immature, taken with the holotype (CAS, examined). 278

Description.—Male. Color brownish. Carapace with three pairs of dorsal and three apical setae. Eyespots indistinct. Anterior sternum with 13 bifid setae. Abdominal terga I-V with two setae, terga VI-IX with four setae, segment XII with acutely produced, well developed posterodorsal process. Vestigial stigmata darker than sterna,

Flagellum pentagonal, with no dorsal relief. Pedipalpal trochanter produced distally, thick spines from tibia appose tarsus-basitarsus, spurs of latter about 1/6, claw about 1/3 length of tarsus-basitarsus.

Tarsal-basitarsal segments of leg I of the following approximate proportions: 51-10-10-10-12-11-24, Other leg segment measurements given in Table 17,

Female, Flagellum composed of four articles, Pedipalpal trochanter not as acutely produced, Spermathecae composed of several pairs of lobes which are apically elaborated with many minute terminal elevations, no special sclerotization.

Comparisons.--This species apparently lacks the high degree of dimorphism of the pedipalps as encountered in other California species with the exception of S^. pentapeltis. The flagellum is completely lacking any dorsal relief, which further contrasts with other briggsi group members. See also under S^. wessoni.

Distribution.--This species is known only from Borrego Palm

Canyon, Anza Borrego State Park, San Diego County, California.

Additional records.--Borrego Palm Canyon, 12 January 1971,

J. Rowland, T. Moisi, two males, one female (AMNH). 279

Schizomus shoshonensis (Briggs and Hom), 1972 NEW COMBINATION

(Figs, 251, 256; Tables 16, 17; Map 6)

Trithyreus shoshonensis Briggs and Hom, 1972:1-7,

Holotype,--An adult male, taken in Upper Shoshone Cave, near

Shoshone, Inyo County, California, on 28 December 1971, by W. Ranscher,

E. Fogarino, T. Briggs (CAS, examined).

Allotype.--An adult female, taken with the holotype (CAS, examined).

Description,--Male, Color brownish. Carapace with one pair of dorsal and three apical setae. Eyespots indistinct, but present.

Anterior sternum with 11 bifid setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segment XII with well developed, acute posterodorsal process. Vestigial stigmata slightly darker than sterna, Flagellum heart shaped, with pair of paramedial elevations separated by a wide median depression which deepens and widens distally, Pedipalpal trochanter produced distally slightly, all segments elongate, tibia with a spur apposible to tarsus-basitarsus, spurs of the latter about 1/6, claw about 2/5 length of tarsus- basitarsus. Tarsal-basitarsal segments of leg I of the following approximate proportions: 48-7-10-9-9-9-23. Other leg segment measurements given in Table 17,

Female. Carapace with three pairs of dorsal setae. Flagellum composed of four articles. Spermathecae not dissected, but appearing typical for the group as seen through the lightly sclerotized genital sternum. 280

Comparisons.--This species is unique among the members of the briggsi group in that it possesses but a single pair of dorsal carapacal setae in the male. Its closest relatives, with which it bears obvious similarities, are more western. Also see under S^, belkini,

Distribution.--This species is known only from Shoshone Cave, near Shoshone, Inyo County, California.

Remarks.--This species is unquestionably troglobitic, as was convincingly pointed out by Briggs and Hom (1972) in their original description. It is likely that other populations of this or related species may occur in surrounding caves. The maintenance of eyespots in this species points out that it may be a fairly recent relict. It also confirms that other species, such as S^. pallidus of the mexicanus group, may also be troglobitic, yet retain some traces of eyespots.

Schizomus joshuensis (Rowland), 1971 NEW COMBINATION

(Figs. 249, 261, 272; Tables 16, 17; Map 6)

Trithyreus joshuensis Rowland, 1971c:304-308; Briggs and Hom, 1972:2;

Rowland, 1972b:3, 4, 5, 7; Rowland, 1972c:153, 155, 156, 159.

Holotype.--An adult male, taken at Forty-nine Palms, Joshua

Tree National Monument, San Bernardino County, California, on 20

February 1970, by J. Rowland and D. Harris (AMNH, examined).

Allotype.--An adult female, taken with the holotype (AMNH, examined). 281

Paratypes.--An adult male and adult female, taken at Forty-nine

Palms, on 22 February 1970, by J, and C, Rowland (AMNH, examined); an adult male and adult female, taken at Forty-nine Palms, on 30

December 1970, by J, Rowland and P, Brashier (AMNH, examined).

Description,--Male. Color brownish. Carapace with three pairs of dorsal and three apical setae, Eyespots indistinct. Anterior sternum with 13 bifid setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segment XII with acutely produced, well developed posterodorsal process. Vestigial stigmata darker than sterna, Flagellum roughly hexagonal, with pair of distinct median depressions flanking slight median ridge, Pedipalpal trochanter, femur, patella, and tibia extremely elongate, the tibia with a mesal spur apposible to tarsus-basitarsus, spurs of the latter about 1/6, claw about 1/4 length of tarsus-basitarsus, Tarsal-basitarsal

segments of leg I of the following approximate proportions: 51-7-9-

9-10-11-25, Other leg segment measurements given in Table 17,

Female,--Flagellum composed of four articles, Spermathecae

composed of four pairs of small, broad lobes, each with minute

terminal elaborations, no special sclerotization.

Comparisons,--This species is usually somewhat larger than

other species in the group, with the exception of S^. pentapeltis.

Details of the male's flagellum serve to distinguish it further from

related forms, all of the briggsi complex. Also see under S^, belkini.

Distribution.--This species is known only from Forty-nine

Palms, Joshua Tree National Monument, San Bernardino County, California, 282

Remarks.--Search of other palm oases of Joshua Tree National

Monument has failed to produce schizomids, but when found should closely resemble S^. joshuensis. The type locality and possibly the only suitable habitat for this species is maintained by a fault-zone spring, which, if caused to go dry, should certainly eliminate this species. Lost Palm Oasis, another similar oasis in Joshua Tree

National Monument, has been drained and made uninhabitable for this and perhaps other interesting relicts.

This species has been collected in relatively low temperatures, where water was crystallized on the soil surface. In such cases I have collected this species on the vertical face of partially exposed rocks.

Additional records.--Forty-nine Palms: 20 February 1970, J.

Rowland, two males, two females, one immature (MCZ); 22 February 1970,

J. Rowland, D. Harris, four females, two immatures (CAS); 30 December

1970, J. Rowland, one female (TTU).

Schizomus briggsi (Rowland), 1972 NEW COMBINATION

(Figs. 250, 253, 262, 268, 269; Tables 16, 17; Map 6)

Trithyreus briggsi Rowland, 1972b:l-9; Rowland, 1972c:156.

Trithyreus belkini, Hom, 1967:216-220,

Holotype,--An adult male, taken on the north face of Rocky

Hill, 2,7 mi, E Exeter, 1500 ft., Tulare County, California, on 28

January 1971 by P. and J. Rowland (AMNH, examined). 283

Allotype,--An adult female, taken with the holotype (AMNH, examined).

Description,--Male, Color greenish brown. Carapace with

three pairs of dorsal and three apical setae, Metapeltidium split,

Eyespots indistinct. Anterior sternum with 13 bifid setae. Abdominal

terga I-VII with two setae, terga VIII-IX with four setae, segment XII

with acutely produced, well developed posterodorsal process. Vestigial

stigmata darker than sterna. Flagellum roughly hexagonal, with vague

median depression followed by two elevations in line. Pedipalpal

trochanter, femur, patella and tibia extremely elongate, the tibia

with a mesal spur apposible to tarsus-basitarsus, the latter with

spurs about 1/6, claw about 1/4 length of tarsus-basitarsus.

Tarsal-basitarsal segments of leg I of the following approximate

proportions: 43-6-6-8-8-9-22. Other leg segment measurements given

in Table 17.

Female, Flagellum composed of four articles, Spermathecae

composed of four pairs of small, broad lobes, each with minute terminal

elevations, with no special sclerotization.

Comparisons.--Of the Californian schizomids, S^, briggsi is most

similar to S^, joshuensis and S. belkini, but is much smaller in nearly

every respect than ^. joshuensis. It is easily distinguished from ^.

belkini by the abbreviation of the convex longitudinal ridge on the

dorsal aspect of the male's flagellum. In S^. belkini the ridge is

broad and extends from the base to the tip of the flagellum, whereas

in S^. briggsi the ridge is narrow and extends from near the middle of 284 the flagellum to the tip, S^, joshuensis also lacks a complete ridge, but differs from S^, briggsi by having two lateral pits at the base of the ridge v/hereas S^, briggsi has none.

Distribution,--This species is known from near Academy, Fresno

County, south to Fountain Springs, Tulare County, California, on scattered rock outcroppings along the west face of the Sierra Nevada foothills.

Remarks.--S^. briggsi represents the northernmost New World schizomid. Its occurence is seasonal, being abundant in winter and early spring, but apparently disappearing as the temperatures increase and humidities decrease. As in S^, joshuensis, and perhaps the other briggsi group species, this species has been found in relatively cold weather. K. Hom (personal communication) reported that he has collected this species under rocks with snow on the surrounding ground. Such temperature adaptations are not characteristic of schizomids in general.

Additional records,--Fresno County: 7 mi, E Academy, 16 April

1967, T, Briggs, one adult (TTU); Squaw Valley, 23 March 1941, S,

Mulaik, four females (AMNH); 7 mi, N Piedra, 21 January 1967, T,

Briggs, one adult (TTU); 1.6 mi. SW Piedra, 21 January 1967, T.

Briggs, one adult (TTU); 1.6 mi. SW Piedra, 21 January 1967, T.

Briggs, A, Jung, W. Lum, V, Lee, G, Leung, M, Wong, K, Hom, 21 adults

(LACM), Tulare County: north face of Rocky Hill, 2.7 mi. E Exeter,

21 January 1971, J. and P. Rowland, 21 adults (TTU); 28 January 1971,

J. and P. Rowland, 40 adults (LACM, AMNH, CAS, TTU); 5 January 1972, 285

J. and P- Rowland, five adults (TTU); north face of Rocky Hill, 2,1 mi,

E town of Rocky Hill, 19 December 1966, T, Briggs, V, Lee, K, Hom, 22

adults (MCZ); northwest face of Rocky Hill, 1,4 mi, E town of Rocky

Hill, 22 January 1967, T, Briggs, A, Jung, W, Lum, K, Hom, 18 adults

(TTU); 12 mi, NE Hammond, 21 March 1941, S, Mulaik, one adult (AMNH);

5 mi, NE Lemoncove, 20 March 1941, S, Mulaik, three adults (AT-INH);

hill, 2 mi, SE Ivanhoe, 18 December 1966, T, Briggs, two adults (CAS);

9 mi. N Woodlake, 22 March 1941, S. Mulaik, three females (AMNH); hill,

3 mi. E Lindsay, 19 December 1966, T, Briggs, V, Lee, K, Hom, ten

adults (MCZ); 6.5 mi, E Fountain Spring, 19 March 1967, T, Briggs,

one adult (TTU); 7 mi. E Fountain Spring, 19 March 1967, P. Lum, V,

Lee, K, Hom, 24 adults (MCZ),

Schizomus belkini (McDonald and Hogue), 1957 NEW COMBINATION

(Figs. 248, 264, 265, 271; Tables 16, 17; Map 6)

Trithyreus belkini McDonald and Hogue, 1957:1-7; Briggs and Hom,

1966:270, 273-274; Hom, 1967:216-220; Rowland, 1971c:304,

308-309; Briggs and Hom, 1972:2; Rowland, 1972b:l, 4, 5, 7,

8; Rowland, 1972c:153, 155, 156, 158, 159.

Holotype.--An adult male, taken at Crater Camp, Santa Monica

Mountains, Los Angeles County, California, on 21 March 1953, in oak

humus, by J, Belkin, R, Schick (AMNH, examined).

Allotype,—An adult female, taken with the holotype (AMNH,

examined). 286

Paratypes.--An adult male, three adult females, taken with the holotype (AMNH, examined).

Description.--Male, Color brownish. Carapace with three pairs of dorsal and three apical setae, Eyespots indistinct. Anterior sternum with 13 bifid setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae, segment XII with acutely produced, well developed posterodorsal process. Vestigial stigmata darker than sterna, Flagellum roughly hexagonal, with a pair of vague median depressions divided by a wide median ridge, Pedipalpal trochanter, femur, patella, and tibia extremely elongate, the tibia with a mesal spur apposible to tarsus-basitarsus, the latter with spurs about 1/6, claw about 1/4 length of tarsus-basitarsus. Tarsal-basitarsal segments of leg I of the following approximate proportions: 42-5-7-7-

7-8-20. Other leg segment measurements given in Table 17,

Female. Flagellum composed of four articles. Spermathecae composed of four pairs of small, broad lobes, each with minute terminal elevations, no special sclerotization.

Comparisons,--This species is generally the smallest of the members of the briggsi group. The details of the male's flagellum are so far the only sure characteristics which distinguish this species from S^. joshuensis and S^. briggsi.

Distribution,--This species is known from the Santa Monica,

Santa Ynez, and San Gabriel Mountains on mainland southern California, and on Santa Cruz Island off its coast. 287

Remarks.--The widely disjunct populations indicate that this species has probably been restricting its range since xerothermic conditions have prevailed in southern California, except for a narrow moist corridor along the coastal foothills, which provides an avenue for northward dispersal. As in other members of this group, the occurance of this species has only been detected during winter and early spring months, apparently retreating in warmer, drier weather into subterranean habitats.

Additional records.--Los Angeles County: Santa Monica

Mountains: 4,7 mi, N Topanga Beach, sycamore litter, 27 December 1966,

A. Jung, K, Hom, two males (MCZ); 7 April 1966, T, Briggs, A, Jung,

K. Hom, one male, one female, one immature (CAS); Topanga Canyon,

21 March 1953, J, Belkin, R, Schick, one male, three females (AMNH);

19 December 1965, T, Briggs, 0, Yang, one female (CAS); 29 March 1952,

R. Schick, one female, two immatures (AMNH); 27 February 1952, J.

Belkin, W. McDonald, two females, six inmatures (AMNH); Malibu Canyon:

Tapia Park, 4 April 1954, L, Moskowski, one immature (AMNH); 7 March

1971, J, Rowland, P. Brashier, two females, one immature (TTU); 16

February 1970, J, Rowland, M. Brand, one male, one female, one immature (CAS); Santa Monica Mountains, April 1953, R, Schick, one female, two immatures (AMNH); San Gabriel Mountains: Eaton Canyon,

28 February 1967, M, Thompson, one immature (LACM); 30 March 1968,

J. Rowland, B. Firstman, two males, three females, one immature

(AMNH). Santa Barbara County: Ose Canyon, Santa Ynez Mountains

(=?0so Canyon, San Rafael Mountains), 26 December 1943, W. S, Ross, Figs, 248-253,—Male's flagella of the briggsi group: 248-252, dorsal views: 248, S^. belkini; 249, S^. joshuensis; 250, S^, briggsi;

251, S^, shoshonensis; 252, S^. borregoensis; 253, lateral view of S^. briggsi, Scale: 1 cm = ,08 mm.

Figs, 254-259,--Parts of schizomids of the briggsi group:

254-258, male's flagella: 254, 255, dorsal views: 254, S_. wessoni;

255, S_. pentapeltis; 256-258, lateral views: 256, S_. shoshonensis;

257, S^. borregoensis; 258, S_, wessoni; 259, female's spermathecae of

S^, pentapeltis. Scale: Figs. 254, 256-258, 1 cm = ,08 mm; Figs. 255,

1 cm = .16 mm; Fig, 259, 1 cm = .04 mm.

Figs, 260-265.--Female's spermathecae of the briggsi group:

260, S^, borregoensis; 261, S^, joshuensis; 262, S^, briggsi; 263,

S^. wessoni; 264, 265, S^, belkini: 264, from the type locality;

265, from Santa Cruz Island. Scale: 1 cm = .04 mm.

Figs, 266-272,--Parts of male schizomids of the briggsi group:

266-269, pedipalps: 266, 267, S^, borregoensis: 266, right, lateral view: 267. right, dorsal view; 268, 269, S^. briggsi: 268, right, lateral view; 269, right, mesal view of tibia and tarsus-basitarsus only; 270-272, lateral view of flagella: 270, S_. pentapeltis; 271,

S.. belkini; 272, S^. joshuensis. Scale: Figs, 266, 257, 269, 1 cm

= ,06 mm; Fig. 268, 1 cm = .12 mm; Fig. 270, 1 cm = ,16 mm; Figs.

271, 272, 1 cm = ,08 mm. 4^^-^^^ Table 17,--Measurements of the members of the briggsi group:

1, three males, S^, pentapeltis; 2, three females, S^. pentapeltis;

3, one male, S^, wessoni; 4, one female, S^, wessoni; 5, two males,

S^, borregoensis; 6, one female, S^, borregoensis; 7, one male, S^, shoshonensis; 8, one female, S^, shoshonensis; 9, four males, S^, joshuensis; 10, four females, S_. joshuensis; 11, five males, S_. briggsi; 12, five females, S^, briggsi; 13, four males, S^, belkini;

14, four females, S^. belkini, 297

TABLE 17 Taxa n 1 2 3 4 5 6 7 1 1,49-1,54 1.42-1.59 1,30 1.24 1.38-1.46 1.40 1.25 2 1,39-1,54 ,52-.57 ,75 - ,59-,62 .43 .62 3 .45-.46 - ,66 - ,61-,62 - .69 4 2.05-2.31 1.41-1.63 1,36 - 1,82-1.83 1.39 1.57 5 2.56-3.28 1.68-1.95 1.60 - 2,25-2,32 1.65 1.85

6 2.04-2.39 1.36-1,57 - - 1.74-1,77 1,22 1.48 7 1.30-1.59 1,08-1,18 - - 1,19-1,30 1,14 1,14

8 1.18-1.37 1,00-1.15 ,96 .96 1,14-1,17 1,00 1,02

9 .69-.78 .55-.67 ,51 .53 ,66-,68 ,56 ,59

10 .81-,93 .68-,72 .64 .64 .84-.87 .66 ,69

11 .69-.78 ,55-.65 .54 .56 .65-.67 .55 .62

12 1.05-1.15 ,90-1,09 .85 ,90 1,03-1,07 .91 .92

13 .51-.59 ,46-.51 .39 ,45 ,51-.52 .46 ,45

14 ,65-.75 .57-,64 .55 ,55 .70-.72 .56 ,57

15 .75-.85 ,64-,70 .59 .60 .71-.74 .61 ,68

16 1.53-1.80 1,33-1,51 1,21 1.31 1.54-1.59 1,34 1.39

17 .74-.79 .65-.73 .55 .60 .69-.70 .65 .61

18 1,15-1,30 .99-1.14 .94 .93 1.18-1.22 1,02 1.01

19 1,01-1.15 .86-1.00 .79 .80 .96-.98 ,83 .91 298

TABLE 17 (continued)

Taxa

n 8 9 10 11 12

1 1.18 1,36-1,48 1,39-1,47 1,22-1,32 1,22-1.40 2 .34 ,63-.65 .38-.44 ,55-.60 .38-.40

3 - .65-.68 - .51-.56 -

4 1,23 1.52-1.75 1.34-1.53 1,22-1.35 1.05-1.25

5 1,42 1.96-2.16 1.60-1.86 1.62-1.70 1,31-1,55

6 1.11 1.50-1,70 1.22-1.45 1,14-1,28 1,00-1,03 7 .99 1,12-1,27 1,01-1,12 .82-1,03 ,82-,98 8 .88 1.02-1.17 ,97-1,10 .72-.92 ,80-,84 9 .51 ,59-,67 .52-.61 ,36-,50 ,45-,52 10 .56 ,69-,77 ,64-,75 .55-,63 .51-,54 n ,50 .58-.69 ,56-,65 .46-.53 .42-.52 12 ,79 .95-1.05 ,89-1,03 .70-.74 ,65-,79 13 .39 ,41-.53 ,40-.46 ,28-,36 .35-,38

14 ,47 .55-,66 ,51-,63 ,41-,46 ,43-,45 15 ,54 ,64-,77 .62-,71 ,46-,57 ,46-,55

16 1,21 1,44-1.56 1.32-1.52 1.18-1,26 1,15-1,30 17 ,57 .64-.73 .59-.70 ,50-,56 ,45-.57 18 ,85 1.02-1,15 ,98-1,09 ,83-,95 ,81-,95 19 ,76 ,94-1,04 ,85-,95 ,70-,82 ,68-.81 299

TABLE 17 (continued)

Taxa

n 13 14

1 1.13-1.25 1.11-1.24 2 .54-.57 .34-,39 3 .45-.52

4 1.14-1.31 1,02-1,13

5 1,27-1,55 1,14-1,30

6 .98-1,13 .90-,98

7 .93-,97 ,82-,87

8 ,79-,88 ,73-,84

9 .44-,53 ,45-,52

10 ,48-.55 ,48-.51

11 ,45-.52 .40-.48

12 .70-.78 .66-.76

13 .34-.38 ,32-.40

14 ,38-,42 ,32-,41

15 ,45-,54 .43-,51

16 1,12-1,25 1.05-1,16

17 ,55-,65 ,51-,60

18 ,77-,85 .73-,80

19 ,70-,78 ,65-.75 300 one female (CAS); Santa Cruz Island, April 1913, one male, one female

(MCZ); Field Station, 19 December 1967, K. Hom, one male, two females, one immature (CAS); Raven's Wood, Canada Del Puerto Canyon, 21

December 1967, T, Briggs, A, Jung, K, Hom, one male, one female, one immature (CAS),

Problem Species

The following four taxa present considerable difficulties in their proper placement and are, therefore, not placed in any of the species groups discussed above,

Schizomus troglobius, new species

(Figs. 273, 276, 280; Table 18)

Holotype.--An adult male, taken in Jackson Bay Cave, Clarendon

Parish, Jamaica, on 21 or 22 December 1972, by S. and J. Peck (AMNH),

Allotype,--An adult female, taken with the holotype (AMNH),

Paratypes,—An adult male (MCZ), two adult females (MCZ, TTU), and five immatures (AMNH), taken with the holotype.

Description,--Male, Color pale brownish. Carapace with three pairs of dorsal, the medians the smallest, and two apical setae.

Eyespots absent. Anterior sternum with 11 bifid setae. Abdoriinal terga I-VII with two setae, VIII-IX with four setae, abdominal segments X-XII slightly elongated, segment XII without evidence of posterodorsal process. Vestigial stigmata darker than sterna.

Flagellum laterally compressed, with complex sculpturing. Pedipalpal 301 trochanter slightly produced distally, tarsal-basitarsal spurs about

1/5, claw about 1/3 length of tarsus-basitarsus. Tarsal-basitarsal segments of leg I of the following approximate proportions: 37-7-10-

7-9-10-22. Other leg segment measurements given in Table 18.

Female, Flagellum composed of three articles. Lateral spermathecae somewhat longer than medians, with sclerotized bulbs, medians with unsclerotized bulbs, laterals curved inwardly.

Comparisons,--This species has only a limited affinity to other

Antillean species. It is the only Antillean species in which the female has a flagellum composed of three articles, but the spermathecal form is like the others of the cubanicus group in having the laterals longer than the medians. The male's flagellum is unique in being laterally compressed, and the male's slight elongation of pygydial segments is further unique among Antillean species.

Distribution,--This species is known only from Jackson Bay Cave,

Clarendon Parish, Jamaica,

Etymology.--The specific name is taken from the Greek troglo- meaning cave, and bios meaning life.

Remarks.--This is the only certain Jamaican troglobite. It may be closer related to the mexicanus group than the cubanicus group on the basis of the flagellum of both sexes, but the male's flagellum is so highly derived that it gives no reliable clue. It may, as in i- armasi, represent a relict of the mexicanus group which inhabited

Carribean land masses before diversification of the cubanicus group. 302

The slight elongation of the pygydial abdominal segments in the males adds further to the confusion of the proper placement of this species,

Schizomus infernalis Rowland, 1975

(Figs, 275, 278, 279; Table 18)

Schizomus infernalis Rowland, 1975:18-20.

Holotype,--An adult male, taken 0,8 km N Ruinas de Palenque,

near Palenque, Chiapas, Mexico, on 25 July 1973, from Berlese samples,

by R. Mitchell and J, Reddell (TTU, examined).

Allotype,--An adult female, taken with the holotype (TTU,

examined),

Paratypes.—An adult male and three adult females, taken with

the holotype (AMNH, examined).

Description,--Male, Color brownish. Carapace with two pairs

of dorsal and two apical setae. Eyespots irregular, but circular.

Anterior sternum with 13 bifid setae. Abdominal tergum I with two

setae, II with four setae, III-VII with two setae, VIII-IX with four

setae, segment XII without evidence of posterodorsal process.

Vestigial stigmata darker than sterna, Flagellum spade shaped, with

a pair of vague dorsal depressions, Pedipalpal trochanter very long,

distinctly produced apically, femur greatly thickened, with one mesal

and two lateral teeth, patella curved downward, expanded distally,

tibia with mesal, subapical curved spur apposible to tarsus-basitarsus,

the latter with spurs about 1/4, claw about 2/5 length of tarsus-

basitarsus, Tarsal-basitarsal segments of leg I of the following 303 approximate proportions: 42-8-9-9-9-10-19. Other leg segment measurements given in Table 18,

Female, Flagellum composed of three articles. Median spermathecae just slightly longer than laterals, the former curved outwardly, the laterals with more distinct terminal bulbs, no special sclerotization.

Comparisons,--This species stands quite by itself in the development of the male's pedipalps. No other species possesses the particularly massive development of the femur and trochanter. The flagellum is like that encountered in the mexicanus group, and has a pair of vague dorsal depressions, but which are present very far forward. Immatures as well as adults are easily distinguished from other Mexican species in having four very strong dorsal setae on abdominal tergum II only. The female's flagellum is also like that encountered in the mexicanus group, but the median and lateral spermathecae are nearly equal in size.

Distribution.--This species is known only from near Ruinas de

Palenque, near Palenque, Chiapas, Mexico.

Remarks,--This species could perhaps be placed tentatively in the mexicanus group, but the highly derived condition of the male's

pedipalps, the dorsal setation of the abdomen, and the unique development of the female's spermathecae obscures its true

relationships. 304

Schizomus sp,

(Fig. 283; Table 18)

Specimens examined,--An adult female and two immatures, taken in the Sierra Nevada, Columbia, by B, Malkin (AMNH).

Description,--Male unknown. Female, Color greenish.

Carapace with three pairs of dorsal and two apical setae. Abdominal terga I-VII with two setae, terga VIII-IX with four setae. Vestigial stigmata lighter than sterna. Flagellum composed of four articles,

Pedipalpal trochanter not produced distally, tarsal-basitarsal spurs about 1/5, claw about 2/5 length of tarsus-basitarsus, Tarsal- basitarsal segments of leg I of the following approximate proportions:

34-5-6-7-7-8-16, Other leg segment measurements given in Table 18,

Spermathecae of aberrant form, perhaps a small median highly convergent unsclerotized pair and a very large lateral highly sclerotized pair.

Comparisons,--The greenish color, four articles of the flagellum, and spermathecal form serve to distinguish this species from other New World species.

Distribution,--This species is known only from the Sierra

Nevada, Columbia.

Remarks.--The combination of characters which serve to distinguish this species so readily from other New World species also confuses its proper cladistic placement within existing groups. It is, of course, conceivable that this species like all those treated 305 separately represents a distinct lineage, divergent from both brasiliensis and simonis South American lineages. In the absence of males no attempt is made to place this species.

Schizomus armasi, new species

(Figs, 277, 281, 284; Table 18)

Holotype,--An adult male, taken at Uvero, El Cobre, Oriente,

Cuba, on 25 May 1972, by L, de Armas (IZC),

Allotype,--An adult female, taken with the holotype (IZC),

Paratypes,--An adult male, an adult female, and an immature, taken with the holotype (Af-INH).

Description,--Male, Color brownish. Carapace with three pairs of dorsal and two apical setae, Metapeltidium split or entire,

Eyespots distinct, round. Anterior sternum with nine bifid setae.

Abdominal terga I-VII with two setae, VIII-IX with four setae, segment XII without evidence of posterodorsal process. Vestigial stigmata darker than sterna. Flagellum spade shaped, with a pair of faint dorsal depressions with slight lateral elevations. Pedipalpal trochanter not produced distally, femur, patella, and tibia elongate, the latter without spurs, tarsal-basitarsal spurs about 1/8, claw about 1/3 length of tarsus-basitarsus. Tarsal-basitarsal segments of leg I of the following approximate proportions: 25-5-6-6-5-6-13.

Other leg segment measurements given in Table 18, 306

Female. Flagellum composed of three articles. Median and lateral spermathecae joined basally, wide, not expanded distally, slightly divergent outwardly.

Comparisons,--This Cuban species is unquestionably most closely related to S^. rowlandi, a species not seen by me. The excellent description and figures given by Dumitresco (1973), however, provide enough evidence to associate these species. The male's flagellum is laterally angular in _S. armasi and also bears a pair of weak dorsal depressions. This structure in S^, rowlandi is gradually curved laterally and is apparently without dorsal relief. A much closer resemblance is obtained in comparison of the spermathecae in which the median and lateral pairs are joined basally, A distinction between the spermathecae of the two species is not offered here. The species are probably readily distinguished in any stage by the troglobite faces apparently acutely developed in _S, rowlandi, S_. armasi is an epigean form, with apparently normal pigmentation. The male's pedipalps are apparently short in S^. rowlandi, but are longer in the only examples of S^. armasi available. The occasional split condition of the metapeltidium in S^, armasi will also help distinguish the latter species.

Distribution.--This species is known only from Uvero, El Cobre,

Oriente, Cuba,

Etymology,--The specific name is a patronym given for Dr, Luis

F, de Armas, the discoverer of this and other Cuban schizomids. 307

Remarks,--It may be desirable to place this species with S^. rowlandi into a small, but geographically and morphologically distinct group, I have not done so here because of the paucity of material available to me. These species may represent ancient dichotomies from a proto-mexicanus group lineage, which may have inhabited

Carribean land masses and which may have given rise to a divergent line now largely extinct, having been replaced by cubanicus group species. Figs. 273-279.—Parts of male schizomids: 212-21B,, flagella:

273-275, dorsal views: 273, S^. troglobius; 274, S^, armasi; 275,

S^. infernalis; 276-278, lateral views: 276, S^. troglobius; 277,

§.• armasi; 278, S^, infernalis; 279, lateral view of right pedipalp of S. infernalis. Scale: 1 cm = ,06 mm. 309

277 278 Figs, 280-284, —Parts of schizomids: 280-283, female's spermthecae::

280, _S. troglobius; 281, S^, armasi; 282, S^. infernalis; 283, Schizomus sp. from Sierra Nevada, Columbia; 284, lateral view of male's right pedipalp of S^. armasi. Scale: Figs. 280-283, 1 cm = ,04 mm; Fig, 284,

1 cm = .12 mm. 311

280

282

283 ii

Table 18,--Measurements of the species of Schizomus not assigned to groups: 1, two males, S^, infernalis; 2, two females,

§.. infernalis; 3, one female, Schizomus sp, from Sierra Nevada,

Columbia; 4, two males, S^, troglobius; 5, three females, S^, troglobius; 6, two males, S. armasi; 7, three females, S^. armasi. 313

TABLE 18

Taxa n 1 2 3 4 5

1 1.14-1.18 1,13-1,17 1,17 1,05-1,07 1,19-1,24

2 .40-.41 .25-.27 ,31 ,32-,33 .29-,29

3 .23-.23 - - ,16-.16 -

4 1.05-1.07 .86-.94 1.08 1.42-1.51 1.39-1,43

5 1.16-1.30 1,04-1,15 1.26 1.78-1.86 1.64-1.71

6 .95-.95 ,78-,82 .93 1.31-1.35 1.27-1.29

7 .78-.82 ,70-,73 .83 1,01-1,02 .98-1.00

8 .74-.82 .63-,69 ,76 .91-,93 ,95-,98

9 .43-.55 ,35-.41 ,45 ,47-,48 ,47-.50

10 .44-.50 ,41-,43 ,44 ,63-,63 ,62-,65

11 .41-.43 .35-.35 .45 ,54-,55 ,52-,55

12 .65-.69 .58-.60 .67 ,79-,81 ,77-,83

13 ,29-,35 ,25-,27 .31 .35-.35 ,36-,40

14 ,27-.35 ,29-,30 .34 ,48-,49 ,51-,53

15 ,42-,42 ,34-.38 ,46 ,39-,60 ,57-,58

16 1,04-1.04 .92-.95 1,11 1,30-1.32 1,32-1.34

17 .51-.54 .41-.46 ,50 ,51-.53 ,52-,57

18 ,69-.72 ,60-,65 ,76 ,89-,89 ,94-.96

19 ,60-,65 ,50-,59 .67 ,80-,82 .80-,83 314

TABLE 18 (continued)

Taxa n f 7

1 ,85--.87 .84--.86

2 ,27--.27 .19- .21

3 ,22--.22

4 ,80- .85 .68- .73

5 ,96- 1.03 .80- .85

6 ,72- ,75 .58- .62

7 ,63- ,65 .57- .58

8 ,54- .55 .48- .52

9 ,30- .33 .26- .30

10 .32- ,35 .29- ,31

11 .32- .32 .27- ,29

12 ,40- .47 .43- ,45

13 ,21- .22 .21- ,22

14 ,25- .27 .22- ,24

15 ,31- .32 ,28- ,29

16 .72- .85 .70- .76

17 .34- .35 .33- 35

18 .52- 53 ,48- 50

19 .46- 61 .41-. 43 CHAPTER IV

PHYLOGENETICS

Introduction

The techniques which I have employed in developing classifications borrow from methods and philosophies developed by cladists (Hennig, 1966), the evolutionary school (Darlington, 1970;

Mayr, 1969), and pheneticists (Sneath and Sokal, 1974). Each of these schools has particular weaknesses in developing multipurpose classifications. Difficulties with the biological species concept, non-dichotomous, temporal speciation (evolutionary gradualism), and the concept of overall similarity, among other theoretical problems, have kept the above factions largely static in their applications in biological classification. This is due also in part to the fact that taxonomic classifications are formally tied to Linnaean hierarchies, and as pointed out by Cracraft (1974) Linnaean hierarchies apparently cannot reflect morphological distance or evolutionary rates. ,Although the efforts of taxonomists to create an optimum methodology in producing classifications have failed, they have succeeded to various degrees in developing valuable special-purpose techniques.

I believe, as held by the evolutionary and cladistics schools, that biological classifications should have theoretical bases, and hence predictive power. Evolutionary, historical geographical, and zoogeographical theories are major bases of my classifications, as are relevant genetic and embryological theories underlying decisions

315 316 about homologies. These theoretical bases are fully developed as the holomorphological and chorological methods in Hennig (1966),

In creating my classifications, I have found it most useful to apply formal cladistic methods in determining the branching sequences, or phylogeny, of the taxa involved, I am thus able to

define strictly monophyletic groups and to recognize sister groups.

If necessary, differential rates of evolution within established

clades can then be recognized. Cladistic and patristic classifications are based on different hypotheses, but they are capable of being

intercompatible. Further, cladistic affinities are most often

implied in patristic classifications. Once cladistic relationships

are delineated and recorded, paraphyletic, or potentially paraphyletic,

units can be recognized where needed to reflect patristic

relationships. I have indicated where possible paraphyly is reflected

in classifying.

The cladistic methods used here are those of Hennig (1966),

Camin and Sokal (1965), and Sneath and Sokal (1974), Where phenetic

data were difficult to code, or were rather scarce, the methods of

Hennig were used exclusively. Hennig's methods combine

holomorphological and chorological approaches to recognizing

monophyletic taxa, with the ultimate design (not practiced here) of

setting an absolute chronology in equating appearances of various

lineages.

Numerical cladistics have been found to be the most helpful in

deducing probable phylogenies. Briefly explained numerical cladistics 317 serves to quantify those taxonomic and evolutionary judgments common to traditional and Hennig's methods so that various elements of the classification can be tested for intercompatibility.

The basic assumption underlying coding, and most evolutionary theory as well, is that "character states can be arrayed into some logical order" (Camin and Sokal, 1965). The order of interest, of course, is that from most plesiomorphic to most apomorphic; the transformation series of Hennig. Since contemporary organisms cannot be ancestral to one another, they cannot possess ancestral character states. What apparently is often the case, however, is that organisms with more proximal ancestry will share more apomorphic character states than similar organisms with a more distant ancestry.

It is conjectured, moreover, that some organisms in a particular

lineage may reflect the morphology of the stem taxon of that lineage more closely than other sister groups. It is conceivable that in any

given lineage one branch of every dichotomy will remain morphologically more similar to the stem taxon than to subsequent branch taxa. From

any set of characters used in a polythetic classification in which

transformation sequences can be recognized, relatively plesiomorphic

and apomorphic taxa can be designated.

The numerical cladistic methodologies proposed by Camin and

Sokal (1965) and elaborated by Sneath and Sokal (1974) were used,

where possible to construct cladograms.

The polythetic family, generic, group, and infra-group

classifications and cladistics are based on as many characters as 318 were determined to be useful in phylogenetic analyses. Each character was independently studied in representatives of appropriate taxa and the various states were then described. Next the characters and character states were ordered in a logical sequence and independently studied to determine relative plesiomorphy and apomorphy. In cases where alternatives in the transformation sequence of character states were apparent, each alternative in direction was recorded. Data matrices were constructed which encorporated various plausible interpretations,

Cladograms based independently on each character were drawn and tested against every other character, A compatibility matrix was thus created which indicated the more widely applicable cladograms and probable miscoded data, which were re-evaluated and receded. Having empirically developed an evaluation of the plesiomorphic to apomorphic character states a monothetic method of arriving at a most parsimonious branching sequence was employed. This methodology removes, or furcates, all taxa, one at a time, in an order from most primitive to most advanced, from the data matrix, allowing subtraction of common character states from remaining taxa. The cladogram thus produced can be adjusted for greater parsimony by manually testing various furcation sequences against total numbers of evolutionary steps. The cladogram provided for each taxon represents the most parsimonious set of alternatives in a branching sequence, which is based on the most reliable combination of coded character states. These sequences 319 are arrived at by comparing each alternative transformation series in a stepwise fashion.

Evolutionary studies have shown that character regression as well as similar morphological apomorphy may often be independently achieved in various taxa. In these cases, comparison of provisional cladograms can indicate characters which have been misread and consequently miscoded. Often an incompatible transformation series will initiate a re-evaluation of a character and will allow a recognition of apomorphic details otherwise unnoticed. Of secondary interest to producing parsimonious cladograms of taxa, the empirical methods used here are valuable in deducing transformational directions in characters which are represented in sister groups of broader taxa.

Transitional sequences are often difficult to recognize where several characters have apparently transformed multidirectional ly from a plesiomorphic state, or where they share a highly proximal cladistic relationship. Such character states may have heuristic value in demonstrating probable monophyletic groups, but do not contribute to the construction of the cladistics of superior taxa.

Discussion of the Cladograms

Two cladograms are presented which illustrate alternative branching sequences of the family and generic taxa. The most parsimonious alternative (Fig. 287) is here taken to be the correct one and forms the basis of the hierarchical classification proposed in this study. An inspection of the alternative cladogram (Fig. 286) 320 and the rules of phylogenetic taxonomy will show that the alternative cladogram does not support the proposed classification. It is thus presented here for consideration, but as indicated by cladistic analyses it represents an unlikely history.

The cladogram of the non-briggsi species groups is, like the others, produced by strict rules and does not incorporate subjective opinion beyond character state coding. It is used here to demonstrate certain weaknesses that may not be apparent in a cursory examination.

Two major super-groups seem to be the mexicanus-pecki-goodnightorum

lineage and the cubanicus-simonis-brasiliensis lineage. Strict monophyly can probably correctly be indicated for each of the cubanicus, simonis, and brasiliensis groups. Due to a lesser degree of apomorphy monophyly cannot as certainly be extended to the mexicanus-pecki-goodnightorum lineage. Although the pecki and goodnightorum groups are probably strictly monophyletic, it is not certain whether they diverged from stock that could fall cladistically within or outside the current definition of the mexicanus group. I

have treated the groups as paraphyletic in the sense that certain members of the mexicanus group may in actuality have closer cladistical

relationships with members of the goodnightorum or pecki groups than to

other members of the mexicanus group. This digression from strict monophyly that governs cladistic taxonomy, however, does not affect

the stability of my formal Linnaean hierarchy.

Of the characters used in the polythetic classification of the

groups a few do not lend themselves readily to interpretation of 321 transitional sequence and synapomorphy. These characters were not used in determining the provisional branching sequences, but were added later-

The cladograms of the groups follow the same rules of formulation and need little explanation. Some confusion may be avoided by understanding, however, what is implied in multiple furcations from a single point, A trifurcation in a branching sequence is not meant to imply simultaneous trichotomy of a hypothetical ancestral lineage, but rather indicates a lack of evidence which would resolve the actual sequence of the pair of dichotomies leading to contemporary taxa. It is particularly obvious

in the brasiliensis group that while basic cladistic information-is

discernable, the details of most species dichotomies are not apparent.

Multiple furcations are indicated in the cladograms of the simonis,

brasiliensis, and mexicanus groups. Cladograms were not devised for

the pecki and goodnightorum groups due to lack of adequate material

upon which to base judgments about character states.

Discussion of the Characters

The following are some of the characters used in the taxonomic

and phylogenetic analyses which warrent discussion. I have tried to

present my ideas underlying subjective evaluation of character states

and to discuss possible sources of error. I have also given my

opinion of the relative value of certain characters at various levels

of taxonomy. Characters not explained in the following text were 322 judged not to need discussion and will probably be familiar to most students of the arthropods,

Carapacal setae. Fig, 3 illustrates the general position of the apical and dorsal carapacal setae in schizomines. The number of pairs of dorsal setae is given for each species and sometimes relative size and placement of the setae are described. The apical setae are not indicated in pairs, but in total numbers.

It appears probable that the ancestral lineages of middle

American schizomines had two or three pairs of dorsal setae; in the latter case, the middle pair may have been reduced in size. It has been my observation that many, if not a preponderance of species, have three pairs of dorsal setae. From this it may be deduced that the schizomine stock that inhabited middle America and gave rise to the extant fauna underwent a reduction in size of the middle pair of setae. After dichotomy of several lineages the middle pair continued diminution in some and perhaps regressed to a larger size in others,

Schizomines with two pairs of dorsal setae arrived at that number by losing the middle pair. It is not certain what the homologies are when four pairs are present. This character has been described in members of the Protoschizomidae but was not used in phylogenetic analyses,

Eyespots, These supposed light sensory devices, present only in the Schizominae, appear to be very simple, although the histological details have not been studied. They are not morphologically comparable to lateral eyes of any other arachnid. 323 and their developmental relationships to the latter may be remote.

Simply described, they are irregular, oval, or triangular pale areas present anteriorly on the carapace of members of the Schizominae,

That these eyespots are primitively absent in schizomidans but represented de novo in the schizomines may not be a popular inference.

However, judging by the simple nature of these structures in the schizomines and their complete absence in the protoschizomids and megaschizomines, I am led to believe it. Regression of optic senses in schizomidans is not seriously questionable; however, the state of regression is common to modern schizomidans in ancient lineages,

I have seen schizomids with a definite local convexity of the eyespots which have also been mentioned by Thorell (1889) and Hansen and Sorensen (1905), Schizomus cambridgii and an undescribed Sumatran schizomid are such species. These structures are only remotely similar to the ommatidial structures present in other arachnids, and

I consider them to be apomorphic rather than plesiomorphic.

Metapeltidium, The states of this character have been used since Thorell (1888) to distinguish genera. Some question as to the significance of the split or entire metapeltidium in monothetically distinguishing genera was first implied by Hansen and Sorensen (1905),

They recommended that states of this character may be of only subgeneric value. This character is shown herein to vary within monophyletic taxa and as such should only be used polythetically.

The division of the metapeltidium into separate plates is variable.

The plates are totally separated by a wide margin in some and may be 324 only partially separated by a narrow basal suture in others. Of the

American Schizominae only members of the briggsi and brasiliensis groups are known to have split metapeltidia, but some members of the latter group have an entire metapeltidium. Species within both families are known to show this character in both, and even intermediate, states.

Anterior sternal setae. The number of setae on the anterior prosoma sternum is always described for each species. The ways in which the number can vary, however, is relatively great and leads to serious difficulties in cladistical coding. This limits the use of the character in phylogenetic studies. Where the numbers of setae were used in cladistics, the homologies of the setal combinations were clearly identifiable.

Abdominal dorsoventral muscles. An important difference between the two families of schizomidans exists in the number of abdominal dorsoventral muscles. The protoschizomids possess, in common with members of the order Uropygida, eight pairs of these muscles. They unite the terga and sterna of abdominal segments I through VIII. There is no reason to suspect that this similarity between uropygids and protoschizomids is coincidental. The cornon ancestors of uropygids and schizomidans are here assumed to have had eight dorsoventral muscle pairs, as are the common ancestors of protoschizomids and schizomids.

The members of Schizomidae do not have the pair of dorsoventral muscles uniting the eighth abdominal terga and sterna. It is possible 325 that these muscles were lost in response to an increase in size or position, or both, of the anal glands in the schizomids. In many specimens of this family the anal glands can be seen (owing largely to the mode of preservation) to occupy nearly all of the internal surface of the eighth abdominal sternum. From these specimens, at least, it is apparent that there is some functional relationship between development of the anal glands and absence of the muscles in question. All schizomidans, as far as is known, have functional anal glands; however, dissection of protoschizomids failed to reveal the physical relationship of the eighth pair of dorsoventral muscles and the anal glands.

Abdominal attenuation. All male members of the simonis and goodnightorum groups and one member of the briggsi group have an attenuation of distal abdominal segments. This character state seems to have been achieved independently in the previous groups, and is also represented in some Old World forms. Abdominal attenuation is most highly developed in S^. goodnightorum, but the abdomen is nearly as elongate in a few other species. The elongation may involve segments V-XII or may only involve the pygydial segments, X-XII.

A telescopic intercalation of the abdominal segments occurs in species with a highly attenuated abdomen, which may be capable of some interflexion. S^. pentapeltis carries, at least at times, the distal parts of the abdomen and flagellum up over the basal parts of the abdomen. All species possessing attenuated abdomen have elongate flagella and short, non-dimorphic pedipalps. 326

Spermathecae, The use of the spermathecae was originally introduced as a taxonomic character by Brignoli (1973), although

Borner (1904) first reported its presence. In Rowland (1973c) I registered my doubt that Brignoli had contributed very much by describing two new species from a vast Mexican fauna on the basis of a new and unstudied character. Among my objections I repeat that the spermathecal form is probably too variable intraspecifically to base formal names upon these structures alone, Brignoli (1974) subsequently took steps to show that the spermathecae could be just as useful as the male's flagellum in distinguishing species. He finally provided sketches of three more species to give some indication of the interspecific variation of the character. Sometime before Brignoli's (1973) original work I had studied the spermathecae,

I decided to postpone use of this character since I found too great an intraspecific variability in samples I used. It was (Rowland,

1973c) my opinion that "upon a thorough study of considerable comparative material the spermatheca (sic,) can be put to some use,"

I have now completed the necessary studies and have found, not

surprisingly, that this character is useful in systematics, but unfortunately is subject to transfigurations by preservation and mounting, and is, at least in some species, highly variable, I

suspect that because the observed variability is often unilateral,

a great deal of phenotypic plasticity disguises much of the

underlying genetic uniformity governing this character. 327

I have discovered and used certain spermathecal traits which are very useful in grouping higher categories. I have refrained, however, as I hope others do, from describing any formal taxon on the basis of the female alone. My recommendation is that the female's spermathecae should always be described if available, I have found the following spermathecal traits useful: number of pairs, relative size of median and lateral pairs, presence or absence of terminal bulbs, and degree and location of sclerotization.

Technical difficulties presented by the large size and high degree of sclerotization of the genital sternum precluded my describing the spermathecae of protoschizomids,

Posterodorsal abdominal process. This structure is present only in males of some species of the Schizominae, It consists of a posterior emargination of the dorsal side of abdominal segment XII,

It may project out over the base of the flagellum, or may only slightly protrude. In most non-briggsi American species the structure is truncate, but may also be pointed or gently rounded. In the briggsi group it is always pointed. The structure appears in Old World schizomines, but not in protoschizomids or megaschizomines,

Flagellum of the males. The male's flagellum has proven to be the most useful single characteristic in species recognition. Its form is usually highly consistant within populations, and is not subject to spoilage by preservation. The male's flagellum is grasped by the female at the inception of the mating act. The details of its morphology then could conceivably be perceived by the female and may 328 be important in mate selection. This would seem to be most important, of course, in areas of ecological and geographical overlap of two or more species.

I have been reluctant to formally describe species without the use of the male. The flagellum and the other secondary sexual characteristics of the male offer the most reliable characters for placing species within groups.

Flagellum of the females. The articles of the female's flagellum are joined either by segmentation and annulation or by annulation alone. The term segmentation does not, of course, suggest that the articles are derived from metameres, but only that they are separated from each other by a slight membranous region, Annulations, by contrast, appear to represent lines of incomplete fusion of two flagellar articles, Flagella are very frequently broken off at an annulation, which attests to the nature of the joint. There is no articulation between units joined by an annulation; however, it is also doubtful that much muscular articulation occurs between articles in the segmented condition.

It is probable that the segmented condition of the flagellum arose before annulations, since this is one of many characters shared with members of the order Uropygida. Apparently all schizomidans have annulated flagella; however, only the members of the Schizominae have them exclusively. The flagella of protoschizomids and megaschizomines are characterized by the presence of both segmentation and annulations. They bear annulations proximally and segmentations Fig. 285.--Schematic diagram of the probable cladistic relationships of the flagella of schizomidans. Letters a-f are articles hypothesized present in an ancestral condition: 1, the hypothetical ancestral state of the flagellum in which all the articles are separated by membranes; 2, the hypothetical ancestral state in which only two articles are separated by membranes, the remainder of which are separated by annulations; 3, the state found in megaschizomines; 4, the state found in protoschizomids; 5, the hypothetical ancestral state in which segmentation is absent; 6, 7, the state found in schizomines. 330

00 CO

.a .0 0 o 331 distally. There are two annulations in the protoschizomids, three in the megaschizomines, and two or three in the schizomines. The protoschizomids and megaschizomines appear to be plesiomorphic in this respect. It would appear, however, that either the elements of the flagella of these two taxa are non-homologous, or that the protoschizomids have undergone a secondary consolidation of proximal elements, leaving no trace even of annulations. I tend to believe the latter alternative, because the terminal elements of the flagellum bear certain setational patterns which are constant, and apparently homologous, throughout the order. These attributes suggest that megaschizomines have the most primitive character state, with segmentation and multiple annulations being a condition that probably attained before their dichotomy with the schizomines. Since the dichotomy of the two family lineages, it appears that protoschizomids have maintained the segmentation, but have apparently lost one of the annulations of their ancestors. Schizomines reflect the most advanced state, having lost any evidence of segmentation, but retaining multiple annulations.

A schematic diagram (Fig. 285) best explains my interpretation of the evolution of this structure since the hypothetical stage when schizomidans had six segmented articles, designated by letters a through f. Protoschizomids and schizomines have in common the fusion of flagellar articles b and c in my interpretation. I consider this fusion of elements independent in the two groups, with the unfused state, as seen in megaschizomines, the common ancestral condition. 332

Cheliceral teeth. The teeth of the fixed cheliceral digit show an advancement in states among the family group taxa. The development of the teeth is not only exceptionally well-contrasted from primitive to derived forms, but is also displayed in intermediate states within the order.

The teeth of the protoschizomids consist of two jutting spikes which protrude at about a 45° angle from the anteroventral margin of the chelicera. There is little doubt that protoschizomids represent a state similar to the first stages of development of the cheliceral teeth of the schizomids.

Teeth of megaschizomines are advanced as compared with those of protoschizomids. Whereas the protoschizomids' basal tooth juts out from the cheliceral margin at a considerable angle, the basal tooth in schizomids is a gradually upward curving process. Megaschizomines have three cheliceral teeth, the uppermost and basal ones probably being homologous to the two teeth of protoschizomids. Since divergence from the protoschizomids, the schizomids attained a curved basal tooth. Schizomids also gained one intermediate tooth in megaschizomines and three to seven intermediate teeth in schizomines.

The uppermost cheliceral tooth in schizomines more often than not bears a slight to well developed cleft, which in some species is developed into two teeth,

Cheliceral serrula, brush and file. The movable finger of the chelicera in species of the Schizomidae has a very distinctive structure on the mesal, submarginal surface which I follow Lawrence 333

(1969) in calling the internal cheliceral serrula, I imply, however, no homology of this structure to structures of the same name in other arachnids. It is composed of many, closely situated, similar teeth.

The function of the serrula is not clearly defined. It is assumed, however, to function in ingestion, or grooming, or both. Members of the two schizomid subfamilies appear similar in the nature and development of the serrula. Protoschizomids do not have a serrula as such, but have a series of rather poorly defined knobs. These are no doubt the homolog, if not very similar to the first stage in the development, of the serrula in schizomids.

Another important cheliceral characteristic that appears to

have arisen with the cheliceral serrula in schizomids is the cheliceral

brush, I use this name because the structure consists of an arc of

usually eight closely situated, similar setae. Whereas the setae

are in themselves not fundamentally different from other feathered

cheliceral setae found in both families, together they form a

structure on the mesal surface of the fixed cheliceral digit which is

unique to the members of the Schizomidae.

Some schizomids have on the ventral margin of the movable

cheliceral finger a series of blunt knobs or teeth which constitute

what I call the cheliceral file. It is totally without evidence in

the protoschizomids. Many schizomines show a reduction in this

character to the extent that a single, obtuse, poorly defined file

tooth remains. Perhaps the primitive state of no file, as in

protoschizomids, advanced to the presence of a well developed file. 334 as in megaschizomines and some schizomines, and then to the final derived state of near absence in some schizomines. It is assumed that early schizomidans had only the primitive protoschizomid type serrula, the file appearing in schizomids only after dichotomy of the two family lineages.

It is conceivable, but highly unlikely, that both the serrula and file were present in ancient schizomidans, the file having become lost in some schizomines and the protoschizomids, the serrula having become lost in protoschizomids, and both having been maintained in the Megaschizominae.

The fact that some schizomines lack a cheliceral file adds an element of uncertainty in coding the character states, I have alternatively coded absence as primitive in Agastoschizomus and

Protoschizomus, but occasional absence as apomorphic in Schizomus and presence in Megaschizomus as intermediate. This assumes that the character has regressed in certain members of Schizomus, The alternative coding would be that absence is primitive in all taxa and that some members of Schizomus and all of Megaschizomus have acquired this character state independently,

Pedipalps, The pedipalps of many species of schizomines are sexually dimorphic. The female's pedipalps of schizomines are all fairly similar, the males being more variable. The trochanter may or may not be produced apically and may even be extremely elongate.

The femur and patella may either be short or may be elongate. These segments often bear an armature of spines in members of the 335 brasiliensis group. The elongation of the femur and patella may even be so great in the male that the pedipalps are carried over the carapace. The tibia is often elongate, and sometimes bears a spur which is apposible to the tarsus-basitarsus. The spur occurs in varying degrees of development. Sometimes several stout spines occur where the spur is found in closely related species. The tarsus- basitarsus bears two subapical spurs, one medial and one lateral.

They are usually very similar in size, but are asymmetrically placed in all schizomines, the medial spur being more proximally placed.

The claw of the pedipalp attaches apically to the tarsus-basitarsus and curves gently downward. Simple, perhaps slightly dimorphic, pedipalps are thought to be primitive, while thickening, great elongation, spurs and other strong dimorphisms in the males are thought to be apomorphic.

The sexual dimorphism expressed in the pedipalps of some species is usually inconsistant. It is usually discovered, once adequate collections are made, that the male's pedipalps can be quite variable. Most males may have highly dimorphic pedipalps,

but a few may have pedipalps like the females of that species.

Basitarsal pedipalpal spurs. Although difficult to quantify,

dichotomies at the generic and family level exist in the development

and placement of the basitarsal pedipalpal spurs. All schizomidans

possess two of these claws: one mesal and one lateral, Protoschizomids

have large, symmetrical spurs, Megaschizomines have smaller, slightly 336 asymmetrical spurs, and schizomines have even smaller and more asymmetrical spurs.

The size of the spur is difficult to interpret in a phylogenetic sense since some degree of dichotomy appears in the

Protoschizomidae, The symmetry of placement and similarity in shape of these tv/o spurs seems more indicative of kinship. It is here supposed that the common ancestors of schizomids and protoschizomids

had symmetrically placed, similar spurs. Their size was probably

large. The symmetry and similarity of the spurs is also apparent

in the megaschizomines, but has some symmetry lost, the medial spur

being slightly more proximal than the lateral. In schizomines the medial spur is always distinctly smaller and considerably more

proximal than the lateral spur. The functional significance of this

specialization is not obvious to me,

Pedipalpal spines, Lawrence (1969) first introduced the use

of "true spines" on the pedipalp as one of several characters helpful

in distinguishing the members of two genera.

The differences between "true spines" and ordinary setae are

vague. I think they are homologous structures and that intermediate

conditions can be expected. His "true spines" can best be

characterized as setae that are extremely thickened, highly

sclerotized, rigidly fastened to the exoskeleton, and which may be

raised on a cuticular protuberance. They probably function in the

raptorial use of the pedipalps. 337

Protoschizomids possess these structures on the patella and tibia and Megaschizomines possess them on the femur, patella, and tibia. No schizomines that I have examined attain the development of these spines as found in the latter taxa, but several species approach the development such that homology is apparent.

Ancient schizomids probably had these spines on the femur, patella, and tibia as do modern megaschizomines. They are reduced to absence on the femur in protoschizomids and are altogether wanting or reduced to setae in many schizomines. The nature of the difference between setae and "true spines" suggests that change from one condition to the other, and even back and forth, could be accomplished relatively easily. The use of this character in a phylogenetic classification must be limited, since direction of the transformation series may be difficult to determine.

Trochanter and femur of the fourth legs. Width to length ratios of the trochanter and femur of the fourth legs vary a great deal among the genera, Megaschizomines and protoschizomids are characterized by a relatively long trochanter and femur, though the latter least obtains in the genus Protoschizomus.

The greatly expanded fourth femur of schizomines is associated with the relative development of the contained musculature. Other

schizomidans lack such acute development, though members of

Protoschizomus approach schizomines in this character. Schizomines

alone are characterized by relatively short and wide fourth trochanters. 338

The extreme attenuation of the fourth femur in Agastoschizomus may relate to secondary specializations to their cave environment.

Color. I have been very imprecise in description of the color of various schizomidans. Only two designations with admittedly little objective justification have been made. Each species has either been described as brownish or greenish. I can imagine others choosing to describe what I have generalized as brown anything from red to yellow.

Less confusion is probably met with in my designation of green species.

It is also true that various parts of the body may be noticably different in color. The chelicerae are often a darker reddish color than the rest of the body. I have arbitrarily chosen to describe the color of the carapace and dorsal abdomen of the abdomen, which corresponds to the greatest part of the external surface.

Remarks

In an earlier work (Rowland, 1973b) I devised a classification of the then recognized genera into what seemed an appropriate system of homogeneous groups. The major contribution offered was the union of Agastoschizomus and Megaschizomus into the Megaschizominae as set apart from the members of Schizominae. As my studies progressed more important differences and similarities began to surface. The first classification now seems inappropriate, even misleading in a phylogenetic sense. Knowledge of the groups, furthered by a critical examination of more material, is now such that a more definitive 339 statement on the probable phylogenetic (cladistic) relationships of these groups can be made.

The characters shared by Agastoschizomus and Megaschizomus, which were the basis for the creation of the subfamily Megaschizominae, were not misrepresented in the early classification; they cannot, in fact, be elaborated upon after much more study. Rather, it was the ignorance of several rather fundamental similarities that Megaschizomus, in contrast to Agastoschizomus, shares with members of the Schizominae that led to the previous classification.

The more recent classification (Rowland, 1975), in essence, allies the Megaschizominae and Schizominae within one family and

Agastoschizomus and its close relative Protoschizomus within another.

Protoschizomidae was created for the latter two genera, and Schizomidae contains the nominate subfamily and Megaschizominae. I have conceived the previous taxa to be monophyletic in the rigid sense of Hennig

(1966).

Relative to schizomids, protoschizomids have apparently undergone less specialization since their divergence. This is pointed up by the fact that they present few character states which can unequivocably be judged as apomorphic when compared to character states in the schizomids.

The megaschizomines share several primitive character states with protoschizomids, and fewer advanced character states with the schizomines. Further, there seems to be little convergence to confuse this classification. 340

It seems only possible, according to the characters used there, that the megaschizomines share a more proximal cladistic relationship with the schizomines than to protoschizomids. To deny this relationship one suggests that all those derived character states shared by schizomines and megaschizomines are independently derived,

Taxonomic relationships of the genus Schizomus to other extant taxa of equal or higher rank were discussed earlier. It is not possible at this point to describe in detail a hypothetical taxonomic unit, in the sense of Farris (1970), ancestral to all members of

Schizomus, Even to judge the relationships of briggsi group species with other American species would be premature, I am only able, with any certainty, to briefly outline the character states of a hypothetical taxonomic unit ancestral to non-briggsi group American species. This

HTU can be characterized as follows: color brownish; carapace with indistinct oval eyespots with three pairs of dorsal paramedial carapacal setae, the middle pair being somewhat smaller than the other pairs; carapacal length about 1.00 mm; metapeltidium entire, free of setae; anterior sternum with about 9 setae; abdominal terga

I-VII with 1 pair of dorsal setae, VIII-IX with two pairs; posterodorsal abdominal segments of male's abdomen similar to female's; no posterodorsal process present on XII abdominal segment; female's flagellum with four sections; male's flagellum dorsoventrally compressed, lanceolate, with a pair or a single dorsal depression; female's lateral and median spermathecae narrow, non-divergent, and similar in size and shape; male's pedipalp much longer than female's, 341 thin without apposable, subapical tarsal-basitarsal spur. These characters are, of course, supplemental to those diagnostic for all relevant higher categories. Several apparently synapomorphic character states are judged to have been independently derived, but only a few regressions to a plesiomorphic state have been hypothesized.

These general rules are in keeping with accepted methods for determining clades by empirical methods.

The numerical methods employed elsewhere are useful here even though there is a relative paucity of shared character states. It is most likely that the middle American ancestral stock underwent rapid and multidirectional adaptations in several characters within a relatively short period. The morphological evidence available for taxonomic purposes is extremely useful in delimiting monophyletic groups by recognition of synapomorphic and autapomorphic character states, but is not equally useful in defining cladistic relationships. 342

Table 19,--Characters, character states and assigned codes of the major lineages of Schizomida, Alternative codes which have proven least parsimonious for various character states are included parenthetically.

CHARACTERS STATES, CODES

(1) eyespots absent=0(l); present=l (0).

(2) metasternal setae six=0(l); four=l(0).

(3) abdominal dorsoventral muscles eight pairs=0; seven pairs=l.

(4) female's flagellum segmented=0; segments fused=l.

(5) articles b and c of female's b and c distinct=0; b and c fused=l;

flagel lum b and c independently fused=-l(l).

(6) cheliceral serrula and brush absent=0; present=l,

(7) cheliceral file absent=0; present=l; absent or

present=^2(-l),

(8) intermediate cheliceral teeth none=0; one=l; three to seven=2.

(9) palpal "spines" present on femur, patella and tibia=0; present on patella and

tibia=l; absent=-l(2),

(10) palpal spurs symmetrical=0; slightly asymmetrical 1; highly asymmetrical=2.

(11) trochanter IV length/width 2,2 and higher=0; 1,4 and lower=l.

(12) femur IV length/width 4,1 and higher=0; 2,9 and lower-1. Table 20.--Data matrix of the genera. Codes in parentheses have proven least parsimonious. The compatibility matrix based on the most parsimonious alternatives requires seven extra steps to accomodate all 12 character patterns and requires 18 steps by the monothetic method. The compatibility matrix based on the least parsimonious alternatives requires 67 extra steps to accomodate all 12 character patterns, and requires 23 evolutionary steps by the monothetic method. Explanation of the characters (n) is given in Table 19,

OTU's are as follov/s: 1, Agastoschizomus; 2, Protoschizomus; 3,

Megaschizomus; 4, Schizomus. C, the number of character states;

M, the minimum number of evolutionary steps required to produce the number of character states. 344

TABLE 20

OTU 's n 1 2 3 4 C M

1 0(1) 0(1) 0(1) 1(0) 2

2 1(0) 0(1) 0(1) 0(1) 2

3 0 0 1 1 2

4 0 0 0 1 2 5 1 1 0 -1(1) 3(2) 2(1]

6 0 0 1 1 2

7 0 0 1 2(-l) 3 2

8 0 0 1 2 3 2

9 1 1 0 -1(2) 3 2

10 0 0 1 2 3 2

11 0 0 0 1 2 1

12 0 1 0 1 2 1 Figs, 286-287.--Cladograms of the genera of schizomidans: 286, the cladogram based on the least parsimonious alternatives in coding, requiring 23 evolutionary steps; 287, the cladogram based on the most parsimonious alternatives in coding, requiring 18 evolutionary steps. Characters (small numbers) and OTU's (large numbers) are explained in Tables 19 and 20. 346

286

287 347

Table 21 .—Characters, character states and assigned codes of the non-briggsi American species groups.

CHARACTERS STATES, CODES

(1) dorsal carapacal setae two or three pairs=0; three or four

pairs, usually four=l; three or

four pairs, usually three=-l.

(2) carapacal length up to 1.42=0; up to 1,48=1; up to

1.74=-1,

(3) metapeltidium entire=0; split-1.

(4) distal abdominal segments short=0; attenuated=l,

(5) male's posterodorsal process absent=0; slight to well developed=l

(6) shape male's flagellum ovoid=0; slender=l.

(7) relief male's flagellum simple=0; medium=l; medium to

complex=2.

(8) female's flagellum four sections=0; three sections=l,

(9) length female's flagellum ,38 and below=0; ,47 and above=l,

(10) spermathecae medians equal laterals=0; medians

larger than laterals=l; medians

smaller than laters=-l.

(11) male's pedipalps dimorphic=0; nondimorphic=l,

(12) color brown=0; brown to green=l, 348

Table 22,--Data matrix of the non-briggsi American species groups.

Explanation of characters (n) is given in Table 21, OTU's are as follows: 1, cubanicus group; 2, simonis group; 3, brasiliensis group;

4, mexicanus group; 5, pecki group; 6, goodnightorum group. Explanation of C and M is given in Table 20,

OTU's

n 2 3 4 5 6

1 0 0 •1 0 0 1 3

2 0 0 •1 0 1 0 3

3 0 0 1 0 0 0 1

4 0 0 0 0 1 2

5 1 1 0 0 0 2

6 0 0 0 0 1 2

7 1 2 0 0 0 3

8 0 1 1 1 1 2

9 0 0 0 1 0 2

10 -1 0 1 1 1 3

11 0 0 0 1 1 2

12 1 1 0 0 0 2 Fig. 288,—Cladogram of the non-briggsi groups of American schizomids. Characters (small numbers) and OTU's (large numbers) are explained in Tables 21 and 22, Characters represented by circles numbers were not employed in determining the branching sequences, but were added afterward. 350

288 351

Table 23,--Characters, character states and assigned codes of the members of the cubanicus group.

CHARACTERS STATES, CODES

(1) dorsal carapacal setae two pairs=0; three pairs, medians

smallest=l; three pairs, equal

sized=2,

(2) male's posterodorsal process vaguely developed=0; slightly

rounded=l; well rounded or

truncate=2; pointed=3,

(3) lateral elevations male's absent=0; present=l; secondarily

flagellum lost=2,

(4) median pit male's flagellum single=0; absent=l; double=-l,

(5) male's pedipalp trochanter not produced, length short, width great=0; trochanter

produced, length great, width

great=l; trochanter not produced,

length great, width narrow=l,

(6) color brownish=0; greenish=l,

(7) median spermathecae medium length, unsclerotized=0; medium length, sclerotized bulbs=l;

short, sclerotized=2,

(8) carapacal length .96-1,10=0; 1,16=1; 1,26-1,37=2.

(9) pedipalpal trochanter notch absent=0; present=l; secondarily reduced=2. Table 24,--Data matrix of the members of the cubanicus group.

Explanation of the characters (n) is given in Table 23, OTU's are as f011 ows: 1, S^, cubanicus; 2, S^, cubanicoides; 3, S^. cousinensis;;;;;;;;;,

4, S_, primibiconourus; 5, S^. longipalpus; 6, S^, brevipatellatus; 7,

S^, subiconourus; 8, S^, monensis; 9, S^, desecheo; 10, S^, biconourus;

11, S^. insignis; 12, S_. peckorum; 13, S^. viridis. Explanation :* C and M is given in Table 20, 353

C^JOOCMCMCMr-c^oJCM

tj

CO •— CM CM

CM O O CVI CM CM

CM O O

0^ CO o o r- ,_

cn -— CM ^ r_ O CM 00 >— CM r- r- O O CM O QQ CO

'—-—>— O O 1— O O o vo O O CM I— r- O O CM O I I vn O O I— O I— O O CM O

O O r- O r— O O O

O O I— o o o o

o o o O r— o O O O

o o o O r- O O O

CM 00 «;^ LO vo t^ CO cn Fig. 289,—Cladogram of the members of the cubanicus group. Characters.(small numbers) and OTU's (large numbers) are explained in Tables 23 and 24, !.!trTV7,f:n^^n^r^.,^,,^••^•^^•^^^.•^^:

355 356

Table 25.—Characters, character states and assigned codes of the members of the simonis group.

CHARACTERS STATES, CODES

(1) dorsal carapacal setae three pairs=0; two pairs=l. (2) sternal setae 11=0; 10=1; 12=-1,

(3) male's posterodorsal process round=0; small, truncate=l; large, truncate=2.

(4) lateral elevations male's present=0; absent=l, flagellum (5) median pit male's flagellum double=0; single=l.

(6) eyespots indistinct=0; distinct=l.

(7) elongation male's abdomen VII-XII=0; V-XII=1; X-XII=-1,

(8) length female's flagellum .37-,41=0; .52-1; .61=2,

(9) spermathecal synmetry medians equal laterals=0; laterals longer=l.

(10) spermathecal shape thin=0; broad=l. 357

Table 26.--Data matrix of the members of the simonis group.

Explanation of characters (n) is given in Table 25. OTU's are as follows: 1, OTU #1; 2, OTU #2; 3, S^. drakos; 4, S^. simonis; 5, S^. tobago; 6, S^. acrocaudatus; 7, S^. trinidanus; 8, ^. flavescens;

9, S^. mummai; 10, S^. centralis. Explanation of C and M are given in

Table 20.

OTU 's

n 1 2 3 4 . 5 6 7 8 9 10 c M

1 0 0 0 1 1 1 1 1 1 1 2 1

2 1 0 -1 0 0 1 0 0 0 0 3 2

3 - - 0 1 1 1 1 - 2 2 3 2

4 - - 0 0 1 0 0 - 0 1 2 1

5 - - 0 0 1 0 0 - 1 1 2 1

6 0 0 0 0 0 0 1 0 1 1 2 1

7 - - 1 -1 0 0 -1 - 1 0 3 2

8 - 0 - 1 0 - 0 2 0 0 3 2

9 0 0 - 0 0 - 0 0 1 1 3 2

10 1 1 - 0 0 - 0 0 0 0 2 1 Fig, 290.--Cladogram of the members of the simonis group. Characters (small numbers) and OTU's (large numbers) are explained in Tables 25 and 26. '''rr«rrr'rr>n^.'ff^f„,•lM^..^:JS,v,.^;ff,•f~^-j^^

359

9 10

12 3 6 8

290 360

Table 27,--Characters, character states and assigned codes of the members of the brasiliensis group.

CHARACTERS STATES, CODES

(1) dorsal carapacal setae three pairs=0; four pairs=l,

(2) metapeltidium entire=0; split=l,

(3) sternal setae 9=0; 10=1; 11=2; secondarily 10=3.

(4) color patella I brown or green=0; white=l,

(5) color greenish=0; brownish=l,

(6) male's pedipalpal armature femur and patella unarmed=0; femur and patella armed=l.

(7) male's posterodorsal process slightly developed=0; well developed=l,

(8) spermathecal number two pairs=0; one pair=l,

(9) carapacal length .91-1,11=0; 1.33-1.48=1.

(10) length female's flagellum .20-,30=0; ,38=1; .47=2. Table 28.--Data matrix of the members of the brasiliensis group.

Explanation of the characters (n) is given in Table 27, OTU's are as follows: 1, S^. stewarti; 2, S^, trilobatus; 3, S_, lacandonus; 4, S^, cuenca; 5, S^. sturmi; 6, S^, brasiliensis; 7, OTU #7; 8, OTU =8; 9,

OTU #9; 10, OTU #10; 11, OTU #11; 12, OTU #12; 13, S^, macarensis;

14, S^, cumbalensis; 15, S^, pallipatellatus. Explanation of C and M is given in Table 20. 362

CO CM

CMCM«*CMCMCMCMCMCMCO

Oi— COi— Or— 1— ooo

Or— COr— r— r— r— Oi— CM

O r— 00 O .— I— I

CM r— 1— 00 .— O OOO

O r— 00 O O OOO

c— 1— OO O O o o CO CM CT> O 1— OO O O O i— o

CO •a: =) CO o —r oo o o ooo

O r— oo O O ooo

vo Or— CO O O '— 1—

LO Or— COOr- r— I— Or-

Oi— CMOr— I— r— Ir—

OO OOCMOOOOr— OO

CM OOr— OOOOr— oo

OOOOOOO r O

I— CM00.=:fl.0VDr^00CT>O Fig. 291.—Cladogram of the members of the brasiliensis group. Characters (small numbers) and OTU's (large numbers) are explained in Tables 27 and 28. 364

Table 29.—Characters, character states and assigned codes of the members of the mexicanus group.

CHARACTERS STATES, CODES

(1) dorsal carapacal setae two pairs=0; three pairs=l.

(2) carapacal length ,98-1,23=0; 1,34-1,44=1,

(3) eyespots indistinct=0; distinct=l; absent=-l,

(4) color brownish=0; greenish=l.

(5) pit male's flagellum double=0; single=l; absent=-l,

(6) lateral elevations male's absent=0; present=l.

flagellum

(7) shape male's flagellum round to triangular=0; elongate=l.

(8) length female's flagellum .27-,30=0; ,35-,37=1; .45=2,

(9) male's pedipalp medium length, slightly dimorDnic=0;

extremely long, highly dimorphic

=1; short, not dimorphic=-l,

(10) spermathecal number two pairs=0; three pairs=l; one

pair=-l,

(11) spermathecal length medians equal laterals=0; medians

larger=l; laterals absent=2.

(12) spermathecae lightly sclerotized=0; heavily

sclerotized=l. Table 30.--Data matrix of the members of the mexicanus group.

Explanation of the characters is given in Table 29. OTU's are as follows: 1, OTU #1; 2, OTU #2; 3, S_, mulaiki; 4, S^, bartolo; 5,

S^, lukensi; 6, S^. davisi; 7, S^, reddel Ii; 8, S^. mexicanus; 9, S^, pallidus; 10, S^, portoricensis; 11, OTU #11; 12, S^, moisii; 13, S^,

:ookei; 1^. S^. mitchelli. Exolanation of C and M -^s i^ver. in

Table 20. 367

z: - - CM - CM - CM CM CM CM CM ^—

(-> CM CM OO CM oo CM oo oo oo oo 00 CM

^ , O J^ o _ o r™ ^ o O •" 1

CO ,_ O , o ^ ,_ o ,_ ^_ 1 o O ^~ 1 CM r— O ,_ , o , o o o o ,_ O '— ,_ ,_ o ,_ ,_ 1 1 o o 1 o ^_ O •" O O o o ,_ o o o o o o , ,_ *" CT» - - o o o o - o o o - - o 00 (/) 00 o o o o o o ,— o 1 o r- O OQ

OTU ' r- o o o o o o CM o - 1 1 1

V£> o o o o o o - 1 o 1 ' 1

UD - - o r— o o o CM o 1 1 1 1

«* o o o o o o o r— - 7 1 -

CO o o o o o o 1 1 1 1 - 1

CO o o - o 1 ' o o • o o o

o o o 1 1 1 o o o - 1 ' -

Ci— CM00C0O CM Fig. 292.—Cladogram of the members of the mexicanus group.

Characters (small numbers) and OTU's (large numbers) are explained in Tables 29 and 30, 369

CN

>o 370

Table 31,--Characters, character states and assigned codes of the members of the briggsi group.

CHARACTERS STATES, CODES

(1) dorsal carapacal setae three pairs=0; three or four pairs=l; one pair=-l.

(2) sternal setae 13=0; 14=1; 11=-1.

(3) color brownish=0; greenish=l.

(4) male's pedipalps short, not dimorphic=0; medium length, slightly dimorphic=l;

long, highly dimorDhic=2.

(5) spermathecae several pairs=0; four pairs=i; three pairs=2.

(6) pit male's flagellum absent=0; vague=l; distinct=2.

(7) shape male's flagellum trilobate=0; triangular to pentagonal =1; hexagonal=2; lanceolate=-l. 371

Table 32,--Data matrix of the members of the briggsi group.

Explanation of characters (n) is given in Table 31, OTU's are as follows: 1, S^, pentapeltis; 2, S_. wessoni; 3, S^, borregoensis;

4, S^, shoshonensis; 5, S^, joshuensis; 6, S^, briggsi; 7, S^, belkini

Explanation of C and M are given in Table 20,

OTU's

n 1 2 3 4 5 6 7 C M

1 1 0 0 -1 0 0 0 3 2

2 0 1 0 -1 0 0 0 3 2

3 0 0 0 0 0 1 1 2 1

4 0 1 1 2 2 2 2 3 2

5 0 2 0 - 1 1 1 3 2

6 1 0 0 2 2 1 1 3 2

7 -1 0 1 1 2 2 2 3 2 Fig. 293.—Cladogram of the members of the briggsi group.

Characters (small numbers) and OTU's (large numbers) are explained in Tables 31 and 32, 373 CHAPTER IV

ZOOGEOGRAPHY

Introduction

The historical geographical model of Middle America formulated

by Maldonado-Koerdell (1964) forms the basis for my reconstructions of the schistic events which produced the present distributions of schizomidans in the New World. Specific studies by Darlington (1938,

1957), Whitehead (1972), Peabody and Savage (1958), Heatwole and

McKenzie (1967), Stuart (1957), Axel rod (1957), Woodring (1954), and

Childs and Beebe (1963), though perhaps contradictory in specific applications, also have contributed to my understanding of New World

zoogeographic evolution. Correlation of presumed phylogenetic and

paleogeographic events are central to understanding and developing

specific hypotheses about evolutionary history, I have refrained from

any speculation whatever where I judged that the available evidence,

either of a phylogenetic or paleogeographic nature, was inadequate.

While no probabilistic statement is forthcoming in analyses of this

sort, I have confined my inductions to the most conservative levels

and have left some hypotheses unnoted for lack of better evidence.

The non-political geographic terminology is that of Maldonado-

Koerdell (1964) except that North America includes Mexican land only

to the Isthmus of Tehuantepec and Middle America includes land from

south of the Isthmus of Tehuantepec to the site of the Panamanian

Portal. South America includes only continental land south of the

374 375

Panamian Portal plus Trinidad and Tobago, The Antilles includes all the insular land in the Caribbean area, except for Trinidad and Tobago,

I have purposely used "middle" as an adjective rather than part of a proper noun in the case of "middle America" and should not be confused with "Middle America," which has been used by zoogeographers and geologists alike to designate a variety of faunal, hypothetical paleogeographical, and Recent topographical boundaries.

The genus Schizomus, as here conceived, embraces a number of as yet scarcely studied species, aside from the now better defined

American species groups. An apparent rapid proliferation of species from at least two Old World sources has provided the Americas with a fauna which has gone largely unfragmented by extinction.

Of the known groups defined in America only the briggsi group

seems to represent a non-sister group in the sense of Hennig (1968) and probably constitutes an Old World element with unique evolutionary

tendencies, Vicariance, in the loose sense, obtains in several

characters between the briggsi group and the other American species

groups.

Exclusive of the briggsi group, American schizomidans probably

form a monophyletic group, that is, no American group other than the

briggsi group is cladistically more proximal to a non-American lineage

of equal or higher rank than it is to other American groups.

The question of the ultimate origin of the non-briggsi stem

lineage which gave rise to the present groups is not easily answered.

It is apparent that South America contributed no endemic lineage that 376 did not arise earlier in middle America, The Antilles also apparently received its only fauna from middle America, A middle American origin of the six groups is here postulated with the possible exception of the simonis group. The latter group probably originated in middle

America, but only proliferated, as did the brasiliensis group, upon reaching South America, The descending stock more recently re-entered middle America,

Mexico above the Tehuantepec Portal has elements of three groups: the mexicanus group, which seems to have undergone most of its radiation there; and the goodnightorum and pecki groups, which, while present above the Tehuantepec Portal, are most diverse below it. The

Antilles has two groups: the cubanicus group, which appears to have arisen in middle America and still shares primitive stock with the latter region; and the mexicanus group, which is present by virtue of the recent and wiae dispersal of its Yucatanian parthenogenetic species

S^. portoricensis.

South America is represented by three groups. The most diverse group is the brasiliensis group which appears to have arisen in middle

America, since its primitive members are Mexican and Costa Rican. The simonis group shows no direct link with middle America in its origin, but advanced members of the group are present there. The mexicanus group, again by virtue of parthenogenetic clones of S^. portoricensis,

is also present.

Middle America, in contrast, contains elements of all six of the species groups. Four of these appear to be broadly sympatric in 377

Chiapas, Mexico, and a partially different four are present in Costa

Rica. Disregarding the parthenogenetic clones of S^. portoricensis as being not indicative of typical schizomid dispersal rates and patterns, it is found that Mexico above the Tehuantepec Portal has three groups, the Antilles has one. South America has two, and middle America is still represented by all six. Using this distribution of group taxa as rude numbers clues in the sense of Darlington (1957) it is indicated that these groups probably arose in middle America.

Since South America was adrift for most of the Tertiary, it seems that the first middle American arrivals were not from that continent. Antillean regions were probably devoid of schizomids until

Late Oligocene or Miocene arrival of cubanicus group members.

The most likely direction from which the primitive middle

American stock arrived is from the north. Direct overland connections into middle American and Caribbean land masses existed from North

America in early Tertiary times, soon after closing of the long lasting Balsas Portal and before dissection by the Tehuantepec Canal.

Protoschizomids, the most primitive schizomidans, are known only from Mexico. The intermediate megaschizomines are known only from Africa, and the apomorphic schizomines have a pan-tropical distribution. This suggests that North America might have been the

Mesozoic center of distribution of now extant fauna. A late Mesozoic movement of primitive lineage from the New World into the Old World may then have prefaced the dichotomy of schizomid and protoschizomid lineages. Upon reaching Africa the schizomid stock then perhaps 378 produced the megaschizomine-schizomine dichotomy. Entry of Mesozoic schizomid stock into the New World may have come from Europe or Asia, but, judging from the close relationships of middle American groups, probably not from both. A later, perhaps Miocene, entry of briggsi group ancestors probably did come from Asia,

Extinctions of divergent sister groups of the early Tertiary schizomine ancestors must account for lack of sister groups in North

America, A widespread Tertiary fauna was probably reduced in the

New World to a relatively few surviving lineages, Agastoschizomus may represent an ancient cave-dwelling group isolated in the Sierra

Madre Oriental since Pliocene and its related genus Protoschizomus has survived in epigean habitats in southern and eastern Mexico.

Schizomine representatives probably were limited to Caribbean and middle American land masses after formation of the Tehuantepec Canal in mid-Tertiary,

Miocene calcite deposits in Arizona contain at least two, perhaps three taxa of schizomidans. They are not well enough preserved to show specific affinity to extant species and it is even likely that

Calcitro fisheri and Onychothelyphonus bonneri are two names for the same organism, Calcoschizomus latisternum, while details of its morphology are also unclear, obviously represents a quite divergent line from existing schizomidans. It is unfortunately impossible to see critical characters sufficiently clearly to make any judgment as to even an approximate cladistic placement of this lineage. It is appropriate to say, however, that a morphologically very distinctive 379 lineage of schizomidans existed in Miocene times in North America, but seems not to have left any Recent descendants.

The cubanicus group

(Maps 4, 8)

Arrival of the cubanicus group lineage into the Antilles prefaced a wide diversification of its species. Rapid evolution with concomittant low extinction rates characterizes the movement of cubanicus group elements through the Antillean area. An unreconstructed series of schistic and reuniting topographical events has allowed several of the intermediate types to be isolated perhaps ecologically as well as geographically. In the Antilles extinctions have been relatively low because this fauna may be largely exempt from incoming waves of immigrant forms from the mainland fauna due to the insular geography.

Probably the only non-cubanicus elements occurring in the

Antillean region are S^, troglobius, S^, rowlandi, S^, armasi, and ^, portoricensis. The former three species are of uncertain relationships, but may represent ancient relicts of an early offshoot from lineages which occupied the common Caribbean-Middle American land masses, S^. troglobius is known from a single Jamaican cave where it is certainly troglobitic. The rowlandi-armasi line represent surface and cave sister species of perhaps an early relict.

The nominate complex of the cubanicus group seems without question to contain the most primitive species of the group, if not Map 8,--Evolutionary zoogeography of the cubanicus group.

The dots represent the cubanicus complex, the stripes represent the primibiconourus complex and the hatching represents the viridis complex. 381 382 the most generalized New World schizomids. The geographical dichotomy of the two contained species, while they are perhaps phenetically and phylogenetically more proximal than sister species in other complexes, leads me to believe that their common ancestry dates back possibly to late Miocene. At this time Middle America maintained a land connection to Cuba, but it had existed, according to Maldonado-Koerdell (1968), to Jamaica since Pal eocene. The upper limit on age of dichotomy of these species is set at late Miocene which is the approximate time at which Caribbean seaways finally dissected the Middle American land connections into the Antilles,

The primibiconourus complex is thus far known only from Jamaica and Haiti, but representatives could seemingly be expected from Cuba and Puerto Rico, The four known members of this complex constitute one pair of closely related Jamaican sister species and a pair of vicariant Haitian species. That the history of S^. brevipatellatus and S^, longipalpus and their stem species is Haitian is indicated by their present distribution; however, the probable age of their dichotomy cannot be conjectured. Of closer morphological similarity and probably closer common ancestry are the sister species S^, cousinensis and S_. primibiconourus. These Jamaican species are known only from caves, but show no certain troglobite facies. Their present distribution indicates that they may represent pockets of a once wider distribution held before highly derived members of the viridis complex had appeared in Jamaica or contiguous land masses. Epigean habitats seem now to be dominated by members of the viridis complex and the 383

parthenogenetic S^. portoricensis of Mexican origin. Extensive surface collections have failed to produce any epigean members of the

primibiconourus complex.

The stem taxon of the primibiconourus and cubanicus groups

apparently occupied the Jamaican-Cuban continuity before extension of

the Cayman Trench (Bartlett Trough), which finally divided the two

masses in Pliocene times. Overland communication of faunas between

Cuba and Jamaica since then were necessarily indirect, perhaps via

Hispaniola.

Divergence of the primibiconourus complex stem antedated

proliferation of the viridis complex, but because the latter group has

a wide distribution in the Antillean region, it is thought that both

lineages arose before large scale dissection of the Caribbean land

masses had occurred.

The viridis complex is the most diverse in the group. Its

species are found on all the Greater Antillean Islands with the

questionable exception of Hispaniola, and they even extend into the

Lesser Antilles. The Jamaican representatives are the most derived

members of the group. S^. peckorum seems to be an early isolate of the

highly variable and now widespread S^. viridis. The latter species and

S.- portoricensis form the dominant elements of the Jamaican schizomid

fauna. S^. peckorum now appears to be restricted to caves in adjacent

parishes in north central Jamaica,

The other pair of sister species in the viridis complex is

Puerto Rican, or at least is nominally so. Desecheo and Mona Islands, 384 both located off the west coast of Puerto Rico, harbor apparent endemics which may represent isolates of a once common Hispaniolan-

Puerto Rican fauna. Most closely related to their stem taxon is the most primitive viridis complex member, the Cuban S^. subiconourus,

Cuba also has among its fauna S^, biconourus and S^, negreai, apparently closely related species and which were derived after the divergence of the Puerto Rican species from the viridis complex lineage. This fact either must conceptually broaden the primal geography of pre-biconourus, viridis complex stock to include Cuba, Hispaniola, and Puerto Rico, or it must be considered that parent faunas were

Haitian and that the extant species represent periferal isolates of their centralized ancestor. Of uncertain zoogeographical relations within the viridis complex is S^, insignis. The uncertain placement of this species limits its usefulness in analysis of distributional history of the viridis complex. It is certainly the species furthest removed from the center of distribution of the cubanicus group, and occurs on

Martinique with S^, dispar, an unstudied taxon of possible relations within the viridis complex.

The brasiliensis and simonis groups

(Maps. 2, 7, 9-11)

The most primitive representatives of the brasiliensis group compose the trilobatus complex. These are strictly middle American, but their ancestors are thought to have given rise to the advanced

South American fauna. That the early radiation of this group took Maps 9-11.--Evolutionary zoogeography of the brasiliensis group,

The stripes represent an early offshoot giving rise to the S^, stewarti lineage, the hatching represents a subsequent offshoot giving rise to the S_, trilobatus-S, lacandonus lineage and to the

South American line, the dots represent the brasiliensis complex. 386 387 place in middle America is strongly suggested by morphological and chorological factors.

The trilobatus complex is unified by several plesiomorphic characters, but also is formed by two distinct lineages, S_, stewarti seems to be the most primitive and may stem from a late Miocene invasion of primitive brasiliensis stock into the Sierra Madre del

Sur in southern Mexico, This movement was perhaps correspondent in time to the closure of the Tehuantepec canal, which had existed since early Oligocene (Maldonado-Koerdell, 1964),

The arrival of stewarti stock beyond the Tehuantepec Portal may have been prefaced by an earlier invasion of mexicanus group stock, though the greatest diversity of the latter occurs northward east of the Sierra Madre del Sur, This may in part be responsible for the lack of diversification of stewarti stock since Miocene times. Evolution of the lacondonus-trilobatus lineage in middle

America probably continued without dichotomy of surviving stock until the close of the Panamanian Portal in late Miocene and in early Pliocene (Hershkovitz, 1969), Ancestors of the lacandonus- trilobatus lineage then moved into South America, which at that time was probably unoccupied by schizomidans. There the brasiliensis complex rapidly diversified. Relicts of the pre-South American invasion are S^, lacandonus and S^. trilobatus.

After arrival in South America the brasiliensis lineage has probably developed several separate lines in various areas, but only scattered representatives are available and allow only preliminary 388 statements to be made on their zoogeographic as well as cladistic affinities,

A distinct line arising from the South American brasiliensis complex is the pallipatellatus lineage. It is constituted by four species of the brasiliensis group in which the terminal half of the patella of the first leg is white. This lineage thus has a monothetic basis and may not contain all of the appropriate species with this and correlated characters to constitute a monophyletic assembalge. Its species in Colombia occur on east, northeast, and southwest faces of the Andes, and thus may be expected from other diverse geographic areas. The middle American representative of this lineage, S^. pallipatellatus, is Costa Rican. It represents no substantial dichotomy from other, strictly South American, brasiliensis group members, hence most probably represents a relatively recent extension of the brasiliensis complex into middle

America,

Dating the arrival of the pallipatellatus lineage in middle

America is not facilitated by well documented topographical events.

The time of this entry into middle America was, however. Pliocene or later if it is believed that the earliest date of entry into South

America of any brasiliensis group stock was early Pliocene, or concommitant with closing of the Panamanian Portal system.

Like the cubanicus group in the Antilles, the brasiliensis group has remained largely undiversified in middle America where they probably originated. Upon reaching previously uninhabited territory. 389

however, these groups underwent diverse and apparently rapid

proliferation.

Distribution of the simonis group (Map 5) is limited to middle

America from Costa Rica southeast into northern South America and

then into Trinidad and Tobago, The species are grouped into three

complexes which roughly correspond to different geographical areas.

The distribution of species seems to indicate that greatest

diversity is reached in northern South America and that advanced

forms, members of the simonis complex, have relatively recently

arrived in middle America from South America,

The simonis group seems to be fairly homogeneous without

extremely primitive or advanced contrasting forms. Rather, a few

clues point to the centralis complex being somewhat more advanced

than the drakos complex with the simonis complex being intermediate.

Each has its own specializations, however, and it is difticult to

point to one complex as being more representative of their common

ancestral stock than the others.

The drakos complex is thus far known to occur only in Guyana

but may have representatives in Surinam. Unstudied species inhabiting

Surinam, well within the possible realm of the drakos group, are

S^. gladiator, S^. surinamensis, and £. vanderdrifti, all probably members of that group. The simonis complex occurs only in Venezuela

and islands off its coast. Schizomus dispar could have relations within the latter complex and may represent an extension of this

stock into the Lesser Antilles. Advanced members of the cubanicus 390 group reached Martinique from the north, thus S^. dispar may be derived from either of these stocks. The only hint in the original description, though certainly not conclusive, is that this species is of a brownish color, rather than the greenish color which predmoninates in the viridis complex of the cubanicus group.

The central is complex is represented in Tobago, Panama, and

Costa Rica. It may have entered middle America at about the time that the pallipatellatus lineage of the brasiliensis group began dispersing northwestward from South America. The two complexes, in fact, have reached Costa Rica, but apparently no further into middle

America. Moreover, the morphological and chorological evidence provided by this group leaves little to be said concerning reconstruction of distribution of stem lineages. It is apparent, however, that the simonis group underwent most of its differentiation in northern South America and has arrived relatively recently in middle America, perhaps with members of the brasiliensis group.

The mexicanus, pecki, and goodnightorum groups

(Maps 3-5, 12-14)

The mexicanus group is divisible into five complexes, and is limited, with one exception, to a distribution above the Tehuantepec

Portal. Members of the mexicanus group are to be found most commonly in the tropical and subtropical associations of eastern and southern

Mexico, though a few collections are available from western thorn forests. Species of this group occurring in the seasonal association Maps 12-14.—Evolutionary zoogeography of the mexicanus group.

Map 12, the hatching represents early movement of mexicanus lineage from middle America into Mexico. Map 13, the hatching represents the moisii complex, the dots represents OTU's #1 and #2, the waves represent the mitchel1i complex, and the stripes represent the movements of the mexicanus-portoricensis lineage.

Map 14, the large hatching represents the mexicanus complex, the waves represent the mitchelli complex, the dots represent OTU's #1 and #2, the small hatching represents the moisii complex, and the stripes represent the portoricensis complex. 392 \ • ^ ^^ Y •' -''l:^, r " ^ -"^W^-'•-, 1 I \ MEXICO r

^<1 : rJ r 7 ^X^ i-u^ 1 Iv / " ^^^^^^ /--a.''? r^ "^-J»y''^^^%^- "0^^^^^^ ', [^ iC ^^fe». '"'- ^feCT - V-'"^-1- ^^S>L] / " "^ ( ' ^^ivS ^^^^KV' ^_I (BELIZE ^^^^R^^^ra^ \, ^^^^m. ">-, / ^^**^^^^^^§ ^^^^m,—--^ ,/. r \ ( r 12 \j GUATEMALA ^ > <' ''^V- \/ 1 ^'"1 S^ "^0 ( \ V_j^ 1 '^x, r\.^|

\\. 1 r '~./ ^i cssSt \ /"-^ ) \ "-L; ^^ V -'^ ?'l ,v T -/' ''-'1 /-^-v" 1 ^ ^\ MEXICO -^—" ; "^ -'~'>'-v^- M ] j A^7 v ^^L --'r-' '^; r-""^^-"^" ,*$^$ 1 1 r^ zf f r^-''l f _ ^M i^<^^^ §^^S / N. V ^''-s '•> 1 '^ ^C, i '•/ J • K; f -^

^^ (BELIZE

^~^ ^ \ V \j ,-' ^ 13 GUATEMALA

*/ » / 1 N I r :* V -'\ frV, ,. *^ '.^ CSX f v. MEXICO

7 ^i. -^ ^ r-' ,---• ./I ^^ V •^-' ) V ^ •../• r'^ / : "^V

/ A: — ' -A ^--^,tJ°'T- '"^

1 '. '•• J 1 1 (BEIIZE

14 GUATEMALA ^" ' "• 393 in western Mexico may have more spotty temporal distribution, perhaps becoming active only in relatively mesic climates, as in species of the briggsi group. Volcanic activity in southwestern Mexico in early

Miocene may be partly to blame for the lack of diversity in that area.

The mexicanus group underwent diversification in Mexico north of the Tehuantepec Portal sometime after the latter closed in the

Miocene and its presumed stock reached there from its origins in middle America. Its ancestors probably arrived in Mexico ahead of the brasiliensis stock, which may be largely responsible for the lack of diversification of the latter group in Mexico. A rapid radiation of species from this basic stock occurred within the mexicanus group in Mexico, as it did with the cubanicus group in the Antilles and the brasiliensis group in South America.

OTU's #1 and #2 are put in a complex by themselves. They seem possibly to closely represent the morphology of an ancestral lineage, but males of these species are lacking. They are apparently peripheral in distribution of this group, occurring in southwestern

Mexico in seasonal habitat.

Members of the mexicanus complex occur in eastern Mexico, The eastern species form a morphologically consistant group. They form several close sister species units, the dichotomies of which are probably mainly Pleistocene. The mexicanus complex probably had a distribution at least to southern Texas and probably well into the southern United States in the last interglacial, Distribution of the northwesternmost species are highly restricted, probably 394 representing relictual habitats of a once broader range. S^. mexicanus is the only member of this northeastern assemblage which still has a broad geographical range, occurring in appropriate epigean and cave habitats along an approximate one-hundred mile, north-south line at the foot of the Sierra Madre Oriental. Populations peripheral in latitude and altitude to S^, mexicanus no doubt represent isolates of a once wider ranging stem species. With the development of more xerothermic conditions in northeastern Mexico, most populations of schizomids were eliminated. Those preserved were located near some permanent source of water or in caves. S^. mulaiki is still to be found along the banks of the Rio Grande in Texas where it is preserved by that permanent water source. South of the Rio Grande S^. davisi is to be found along the banks of the San Fernando River in Tamaulipas.

Other surface drainages in this area no doubt harbor relicts. S_, bartolo is a troglobite, limited to a cave in otherwise extremely xeric habitats in Nuevo Leon. Another suspected troglobite, ^, lukensi, inhabits caves in coastal lowland areas where Pleistocene and Recent drying may account for its isolation, S^, reddel Ii presents certain troglobite facies and may represent a fairly young altitudinal relict of a reddelIi-mexicanus stem species.

Two Oaxacan sister species, S^. moisii and OTU #11, form a complex which is distinct from the distributional center of the mexicanus complex. Late Miocene erogenic activity of the Sierra Madre

Occidental and emergence of the Mexican Plateau and Pliocene vulcanism 395 may have been a considerable factor in the isolation of this complex as well as the southwestern representatives of the mexicanus complex.

Another pair of sister species which here constitute a complex is S^. cookei and S^, mi tchel 1 i, These are very closely related troglobites which inhabit geographically proximal caves. Their common ancestry may, in fact, go back to a cave dwelling ancestor.

These species are autapomorphic to an extent that it is not yet possible to ascertain their cladistically closest relatives.

They are apparently representative of a lineage which no doubt shared stem relations within the mexicanus group, but which has suffered high extinction and very local radiation. Perhaps the mitchelli complex was the first mexicanus group species to reach northeastern

Mexico for colonization, but which were overtaken and largely replaced by mexicanus complex members.

S_. mitchelli and S. mexicanus are apparently parapatric at the transition of cave and epigean habitats. S^. mitchelli has not been taken outside, nor S^. mexicanus inside caves in question.

Competitive exclusion may have existed since the pre-Pleiostocene invasion of mexicanus stem species into northeastern Mexico.

A sympatric relationship of S^. cookei and S^. mexicanus exists wherever S.. cookei is found. This probably related to an increased energy flow into S^. cookei caves rather than a drastically different competitive arrangement between it and S^. mexicanus compared to the

latter species and S^, mitchelli. 396

The portoricensis complex is composed of two morphologically very similar, but otherwise unlike, species. S^. pallidus is a probable troglobite, known from a single cave north of the Tehuantepec

Portal in Veracruz. S^. portoricensis is the most widely spread New

World species and occurs in sexual and parthenogenetic populations, i- portoricensis represents a number of disjunct parthenogenetic clones ranging from Bermuda and Florida, south into the Antilles, southern Mexico, and Central America, and further south into South

America and the Galapagos Islands, Also included are sexual populations known from middle America, the ancestors of which gave rise to the parthenogenetic clones,

S^. portoricensis stock no doubt arose, as others of this group, in Mexico above the Tehuantepec Portal. However, it did not proliferate until reaching the Yucatan Peninsula, S^, pallidus represents an early stem dichotomy, reaching its current position as the portoricensis complex lineage moved southeast into Chiapas and

Guatemala and finally into the Peninsula. From the Yucatan Peninsula the parthenogenetic populations could have rafted onto the Greater

Antilles and Florida, Bermuda may have been reached from Florida or directly from Yucatan, Southward and southeastward dispersal through

Central America and finally into South America probably prefaced their ultimate arrival on the Galapagos Islands,

Lineages with certain synapomorphy within the pecki group are the firstmani and pecki complexes. The latter complex is completely contained below the Tehuantepec Portal, This may indicate a dichotomy 397 from the mexicanus lineage which predates the latter's arrival in

Mexico north of the Tehuantepec Portal. More advanced forms may then have arisen in northernmost middle America and moved into North

America proper. Members of the firstmani complex are synapomorphic in several respects and form an assemblage transitional from the pecki lineage. They may have reached to the north of the Tehuantepec

Portal before radiation of its species. After radiation in Mexico the pecki group perhaps re-entered middle America, This group is represented there by OTU #2, The latter species could possibly, however, represent a middle American origin of the complex.

Three of the four species contained in the goodnightorum group are found south of the Tehuantepec Portal while the other is immediately north of it in Veracruz. The group is highly synapomorphic and morphologically very consistant among its species, S^, goodnightorum seems to be divergent in some respects from the rest and certainly occupies the newest geographical area, but whether ancestors of this species entered Yucatan from the east with S^. portoricensis stock or from the south or southwest is uncertain. Too little information on this group is available to make judgments about its zoogeographical history.

The briggsi group

(Maps 6, 15-17)

The evolution of the briggsi group is better indicated by geographical distribution (chorological data) of its species than Maps 15-17.--Evolutionary zoogeography of the briggsi group.

The hatching represents movement of an early pentapeltis-briggsi lineage, the stripes represent the briggsi complex, and the dots represent the pentapeltis complex. 399 400 by pure morphological evidence (holomorphological data). Species of this group are known thus far only from Arizona and California, including islands off the latter's coast. Existence of relict populations in desert oases and the seasonal nature of other populations give evidence, however, that members of this group can be expected to be found in coastal chaparral in Baja California and pockets of suitable habitat known to occur further inland.

The briggsi group, however, seems to be without representation in Mexico. Its species show character states which seem not to be transitional from those encountered in other New World species groups. The stem species of this group is conjectured possibly to have arrived in North America after radiation of a more temperate adapted Asian stock.

Temperate briggsi stock could have moved across the Bering

Land Bridge under Miocene pluvial conditions, to the exclusion of tropically adapted Old World schizomids. After arrival in the New

World, colder interpluvials probably forced ancestors of this group

(perhaps by that time diverged into the two or more complexes) into more southern areas, perhaps as far south as the northern Mexican

Plateau,

Distributional evidence shows that briggsi species probably had been eliminated from central California and northward since

Miocene times when other fossil groups, presumably of tropical origins, occurred in Arizona. Had briggsi species survived in areas above about the 35° parallel, we could expect pre-Pliocene relicts 401

above and to the west of the region of the San Joaquin Embayment.

Interpluvial, late Miocene briggsi ancestors were probably restricted

in range below 35° latitude but re-entered coastal California in late

Pliocene, or at least after the formation of the San Joaquin

Embayment, These elements were probably part of what Axel rod (1957)

termed a Madro-Tertiary fauna and flora moving into the southwest

United States from the Mexican Plateau,

The pentapeltis complex seems to have solely occupied coastal

areas of southern California, perhaps reflecting its earlier isolation

in the Peninsula in an interpluvial, while the briggsi complex stem

species was concomitantly isolated in mainland habitats. Re-entry

into southern California by the briggsi group then came from two

independent directions: the pentapeltis complex from Baja California

and the briggsi complex from the Mexican mainland.

It appears that pentapeltis reached into California along tne

southern coastal mountains before briggsi complex progenitors

invaded. The belkini complex probably never entered Baja California

or coastal California south of Los Angeles due to previous occupation

by pentapeltis. All briggsi group species appear to be allopatric,

though the ranges and habitats of S^. belkini and S^. pentapeltis are

possibly parapatric between Altadena and Pomona in the San Gabriel

Mountains, They may have been sympatric near the coastal base of the

Santa Monica Mountains before urban development of the Los Angeles

basin. 402

The presence of an isolated population of S^, pentapeltis east of the San Jacinto crest indicates a somewhat wider distribution of this species in the past, probably Recent times. An oasis relict of the briggsi complex, S^. borregoensis, indicates, however, that S^. pentapeltis probably never occurred much east of the Peninsular

Mountains since the end of the Wisconsin glaciation.

Apparently the briggsi complex extended from its

Madro-Tertiary origins northwestward to the Pliocene-Pleistocene coastline and to about 37° latitude. Its eastward range extended at least to central Arizona where a relict population of S^. wessoni exists. The ancient coastline of the Pliocene and Pleistocene epochs

(Peabody and Savage, 1958) may explain the distribution of this complex in coastal central California. The San Joaquin Embayment blocked northward and westward movement of briggsi complex stock into areas beyond. Thus schizomids could only have reached the

Santa Lucia Mountains from the north. The climate in the area offering a northern passage into the Santa Lucia Mountains was probably too cold, hence today no schizomid is present in the apparently suitable habitat in the coastal mountains much beyond

Santa Barbara. When the San Joaquin Embayment drained, schizomids conceivably might have been able to reach the Santa Lucia and San

Rafael Mountains by westward overland dispersal. Because of a decrease in precipitation caused by the Sierra Nevada and coastal mountain uplift, conditions in the valley became too dry for habitation very soon after drainage. 403

Schizomids along the coast of the embayment probably reached there by movement over the Sierra Nevada Mountains previous to the

Pliocene and Pleistocene uplift from 3,000 to 8,000 feet (Axelrod,

1957), S^, briggsi is isolated by the high Sierras to the east and by the relatively arid San Joaquin valley to the west and is only

present on scattered rock outcroppings at the base of the Sierras

from Fresno south to Porterville, The foothills become increasingly dry south of Porterville and are unsuitable for dispersal into

southern areas (Rowland, 1972c),

In addition to limiting S^, briggsi from coastal ranges, the

San Joaquin Embayment also formed a barrier to the north along

present coastal routes for S^, belkini, present in what is now the

Santa Monica and San Gabriel Mountains, Subsequent to the drying of

the seaway through the Santa Barbara area and orogeny of the San

Rafael and Santa Monica Mountains, S^. belkini has apparently moved

northward as far as Santa Barbara, where it occurs in the Santa Inez

Mountains.

As early as late Pleistocene and lasting until perhaps

80,000 years before present, a land bridge extended from the Santa

Monica Mountains to the Santa Barbara Channel Islands (Peabody and

Savage, 1958), This enabled S^, belkini ancestors to populate at least

Santa Cruz Island and no doubt the others as well. Subsidence of this

land bridge, of course, isolated the insular from the mainland

populations. 404

At the end of the Wisconsin glaciation xerothermic conditions caused increasing aridity and produced large desert areas east of the

Sierras and southern coast ranges. The Madro-Tertiary elements present in these areas then were either eliminated or preserved in isolated pockets of optimum habitats.

The farthest outlying Californian relict of the briggsi complex is S_. shoshonensis. It is a troglobite, apparently isolated in Shoshone Cave near Death Valley. It is morphologically closer related to S^. briggsi, belkini, and joshuensis than to S_, wessoni and S^, borregoensis. This dichotomy of species within the briggsi complex seems to correspond with the division of California deserts into upper and lower parts. The biotic assemblages present in more mesic times may have been such that progenitors of these two lineages developed widespread, but distinct populations corresponding to the sharply contrasting altitudes. This may account for the geographically more proximal, yet morphologically non-correspondent

S_. borregoensis, a species with stronger morphological affinities with S^, wessoni of central Arizona,

The other highly relictual population closely related to S_, briggsi and S^, belkini is S^, joshuensis, which is found in a single desert-palm oasis. Considering geographical evidence, it most probably had become isolated since S^, shoshonensis, perhaps in the last 30,000 years, and perhaps much after isolation of the S^, belkini population on Santa Cruz Island, 405

The borregoensis-wessoni lineage probably dichotomized during the same xerothermic revolution that isolated S^. shoshonensis, S_. joshuensis, and populations of S^, pentapeltis in now predominantly desert areas. S^. borregoensis is confined to a desert palm oasis and S^. wessoni was described from a single specimen from along the banks of the Santa Cruz River near Tucson, Arizona. Agricultural development of the area has caused extensive and long term drying of the habitat at the type locality. S^. wessoni probably no longer occurs at the type locality and perhaps has been eliminated entirely.

Another apparently relict population occurs at a spring in Organ

Pipe National Monument, Arizona, but is represented by a single,

inmature specimen that offers no more than a split metapeltidium as

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