AMERICAN MUSEUM Norntates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 3056, 32 pp., 27 figures, 4 tables March 23, 1993

A Reanalysis of Mygalomorpl Families (Araneae)

PABLO A. GOLOBOFF1

ABSTRACT

The higher classification of the mygalomorph ina should be relimited to include the is reviewed. A data set of 42 taxa and 71 and Rastelloidina. The group characters is analyzed, applying a new method for Crassitarsae should include also the Microstig- character weighting. Most families were repre- matidae. The group formed by the Diplurinae and sented by two or more genera, so that their mono- Rastelloidina plus Crassitarsae is called Bipectina. phyly could be tested. The trees that best conform Three of the currently recognized families appear to the data have Atypidae plus , as paraphyletic: Cyrtaucheniidae (in terms of Mecicobothriidae, , non-diplurine Domiothelina), (in terms of Micro- "diplurids," and diplurines as successive sister stigmatidae and Theraphosidae plus Paratropidi- groups ofRastelloidina plus Crassitarsae. Neither dae plus ), and (in terms Fomicephalae nor Tuberculotae appear as mono- ofCrassitarsae plus Rastelloidina). Support for the phyletic groups. The groups (restricted monophyly of Hexathelidae is weak; the data are to Atypidae plus Antrodietidae) and Avicularioi- explained almost as well by trees in which the dea (the rest of the ) are resur- family is paraphyletic. rected. The groups Orthopalpae and Quadrithel-

INTRODUCTION The higher classification of the spider in- ra incognita. The mygalomorphs had been fraorder Mygalomorphae was recently ana- traditionally divided into atypoids (atypids, lyzed by Raven (1985). Prior to Raven's work, antrodiaetids, and mecicobothriids, families the relationships among the families were ter- with a complex male palpal bulb, traces of

I Cornell University/American Museum ofNatural History Graduate Training Program in Systematics.

Copyright © American Museum of Natural History 1993 ISSN 0003-0082 / Price $3.40 2 AMERICAN MUSEUM NOVITATES NO. 3056 abdominal segmentation, and six spinnerets) simony analysis. Using these new tools, Ra- vs. non-atypoids (the remaining families, with ven (personal commun.), Coddington (per- male bulb sclerites fused, completely unseg- sonal commun.), and I have now reanalyzed mented abdomen, and only four spinnerets). the relationships among the families of My- Platnick (1977) showed that the atypoids galomorphae using the characters mentioned shared only plesiomorphies, and were thus in Raven (1985). Given Raven's data, if ev- likely to be a paraphyletic group. Raven (1978, ery one of the 15 families he considered is 1979, 1980, 1981, 1982), Raven and Platnick treated as a terminal and scored as having (1981), Gertsch and Platnick (1979, 1980), the character states he considered plesiom- and Platnick and Shadab (1976) tried to re- orphic for each family, Raven's familial tree solve some groups, notably the "Dipluroi- (1985: fig. 1) is one among several most par- dea" (mecicobothriids, microstigmatids, di- simonious trees. This, however, does not plurids, and possibly all or some guarantee optimal results: even ifeach ofthe "pycnothelids"). Finally, Raven (1985) di- 8 cladograms presented by Raven is in itself vided the mygalomorphs into two groups, maximally parsimonious, putting all those Fornicephalae and Tuberculotae, both in- cladograms together may result in a less than cluding some atypoids and some non-aty- optimal solution. In fact, some lines of evi- poids. The interested reader is referred to Ra- dence suggest that some of the families con- ven (1985), and references therein, for a more sidered by Raven (1985) might not be mono- historical perspective. phyletic. The best example is the Nemesiidae; Raven's (1985) paper was the first to at- Raven (1985: 28) discussed the possibility tempt an understanding of the relationships that some of the proposed synapomorphies within the Mygalomorphae as a whole. Con- of Nemesiidae might actually be synapo- sequently, it became a starting point for the morphies of the more inclusive group Cras- higher systematics in Mygalomorphae. Cod- sitarsae (secondarily modified in most non- dington and Levi (1991) rightly referred to nemesiid Crassitarsae). This would leave the Raven's results as "impressive." monophyly ofNemesiidae unsupported, and At the time of Raven's study, however, it would be unwarranted to use "Nemesi- computerized parsimony programs were not idae" as a terminal in the analysis. Other cases readily available, and analysis could be done are discussed below (see under Taxa). only by hand. Raven explicitly attempted to The aim of this paper is to present a rean- minimize homoplasy for his data, which had alysis of the relationships of the Mygalo- nothing less than 240 terminals (genera). For morphae. I intend to show that some of the that many taxa, finding the optimal tree(s) is, hypotheses of monophyly proposed by Ra- because of the extremely large number of ven (1985) are indeed weakly supported or cladograms to be considered, a very difficult counterindicated by the data, and that a dif- task. Raven (1985) divided the mygalo- ferent tree is a better hypothesis of mygalo- morphs into 15 families, proposed a familial morph relationships. This paper also consti- cladogram, and provided separate generic tutes the first application of a new method cladograms for most of the families. The for character weighting. The method is being number of terminals in the cladograms var- described in detail elsewhere (Goloboff, in ied from 7 to 26. It appears very likely, then, prep.), but a short discussion can be found that some of the trees that Raven presented in the section on "Methods." were not actually most parsimonious trees; This paper uses data that have been col- in fact, some cladogram nodes are said to be lected over several years, in the course of supported by characters which are ambigu- studies focused on aspects other than the re- ously, or even incorrectly, optimized. lationships of mygalomorph families. I wish Not that Raven was unaware of this prob- to express my gratitude to a number of ar- lem, of course. In recent years, powerful mi- achnologists who have helped me in the crocomputer parsimony programs have been course of such studies: Maria E. Galiano, developed (see Platnick, 1989; Fitzhugh, Norman I. Platnick, Robert J. Raven, Fred- 1989), the use of which has also increased erick A. Coyle, Martin J. Ramirez, Emilio A. general awareness ofthe complexities ofpar- Maury, Charles Griswold, and Jonathan A. 1 993 GOLOBOFF: REANALYSIS OF MYGALOMORPHS 3

Coddington. Specimens kindly lent by Rob- (Goloboff, in prep.). Given two trees A and ert J. Raven (Queensland Museum, Bris- B, choosing B instead of A may save a step bane), Norman I. Platnick (American Mu- for one character and require an additional seum ofNatural History, New York), Emilio extra step in another. Even when the two trees A. Maury (Museo Argentino de Ciencias Na- require the same number ofsteps, ifthe char- turales, Buenos Aires), Herbert W. Levi (Mu- acter in which we could save some steps shows seum ofComparative Zoology, Harvard), Ana more homoplasy on the trees being com- T. Da Costa (Museu Nacional de Rio de Ja- pared, tree A should be kept. If by choosing neiro), and Jonathan A. Coddington (Smith- B instead we would save a step in a character sonian Institution, Washington) were rele- which shows less homoplasy, B would be a vant for the present study. Maria E. Galiano better choice than A. It is possible to take kindly shared her unpublished observations into account automatically the relative of postembryonic development of several amounts ofhomoplasy ofthe characters when mygalomorphs. I also wish to express my counting steps to compare trees. This can be gratitude to a number of nonarachnologists done by searching for trees in which the "total (nobody is perfect!) who have also helped me: fit" (i.e., the sum of the fits for all the char- James M. Carpenter, James S. Farris, James acters) is maximum, where fit for each char- K. Liebherr, Kevin C. Nixon, and Quentin acter is measured with a nonlinear, concave D. Wheeler. My wife Claudia A. Szumik used function of the homoplasy (see Goloboff, in the computer program developed for this pa- prep., for a full discussion). A well-known per to analyze her own data on embiids, and concave function of the homoplasy is the helped eliminate many errors. Norman I. (character) consistency index, c, ofKluge and Platnick, Robert J. Raven, Frederick A. Coyle, Farris (1969). The effect of using that func- and Maria E. Galiano provided helpful crit- tion to measure fit is that the difference in fit icisms ofdifferent versions ofthe manuscript. between two trees will be smaller for char- Financial support from the American Mu- acters with more extra steps. seum ofNatural History and the Department An MS-DOS computer program, Pee-Wee of Entomology of Cornell University are 1.45 (available from the author), was used to greatly appreciated. calculate the fit for each character using the consistency index (multiplied by 10, with one METHODS significant digit), and to search for trees with Abbreviations. Abbreviations used in this higher total fits. Pee-Wee's branch-swapping study are standard for the group: PLS, pos- algorithms are less effective than those ofoth- terior lateral spinnerets; PMS, posterior me- er parsimony programs (such as Hennig86); dian spinnerets; ALS, anterior lateral spin- to increase the chance of finding all or most nerets; STC, superior tarsal claws; ITC, ofthe optimal trees, several runs were made, inferior tarsal claw. using different initial trees. Parsimony analysis. A data set in which For some of the characters, the terminals 42 taxa were scored for 71 characters was may have more than one state. Those steps analyzed. As pointed out by Coddington and "within" terminals were counted to calculate Levi (1991), cladistic analysis is essentially a the fit for the corresponding characters. For problem of how to allocate homoplasy. If simple step-counting analyses, those steps are there is conflict between two characters, pos- uninformative (they are the same in all the tulating extra steps for one or the other char- trees), but in the present context they are in- acter may not be equally reasonable. It is bet- deed relevant, because they lower the weight ter to postulate homoplasy for those ofcharacters which are known to be variable characters which are more discordant with for some terminals. the tree. This forms the basis for some meth- Hennnig86 version 1.5 (Farris, 1988) was ods of character weighting, such as "succes- used to find shortest trees in the preliminary sive weighting" (Farris, 1969; Carpenter, analyses, to handle the trees produced by Pee- 1988; Platnick et al., 1991). Wee, and to calculate consensus trees. Clados The weighting method used here is derived version 1.2 (Nixon, 1992) was used to find from successive approximations weighting most parsimonious optimizations. 4 AMERICAN MUSEUM NOVITATES NO. 3056

Figs. 1-3. Male palps. 1. Megahexura fulva (Mecicobothriidae), right palp. 2. Stenoterommata pla- tense, left palp. 3. Pseudonemesia sp. (Colombia), right palp.

TAXA present on the highly modified leg I of Cal- ommata, the spines on leg II conform to the Atypidae. Four atypid synapomorphies above description); to a lesser extent, the have been proposed (Gertsch and Platnick, metatarsal trichobothria are displaced pos- 1980; Raven, 1985): maxillae rotated, very teriorly. No other mygalomorph has similar- elongated and curved maxillary lobes, tarsal ly modified spinnerets or legs. Given that claw teeth on a common process, and trun- there are no serious reasons to doubt the cated PMS. The truncation of the PMS re- monophyly of the group as currently consti- sults because the spinning field ofthose spin- tuted, it is included as one terminal. nerets forms a continuous surface with the Antrodiaetidae. Coyle (1971) and Raven spinning field ofthe second article ofthe PLS (1985) considered thatAntrodiaetus plus Aty- (and covers the basal article ofthe PLS, which poides form a monophyletic group, based on does not have spigots) (see Gertsch and Plat- the shared anterodorsal male cheliceral pro- nick, 1980: figs. 23, 49). The labium and ster- cess and reduction of ALS, and possibly the num of atypids are also unique among my- strongly developed inner conductor. No galomorphs; the sternum is abruptly narrowed characters which would be apomorphic for in front, and the labium is much narrower any placement are shared by this group and than the sternum (Gertsch and Platnick, Aliatypus, the only other antrodiaetid 1980). One additional, sixth character is that (provisionally included in Antrodiaetidae by the anterior legs ofatypids are outwardly "ro- Coyle, 1971: 393). Given that the monophyly tated" in such a way that the prolateral spines of the family seems arguable, the two groups have become dorsal (although no spines are were included separately in the analysis. 1 993 GOLOBOFF: REANALYSIS OF MYGALOMORPHS 5

Cyrtaucheniidae. Raven (1985: 61, 65) Platnick (1987) gave two synapomorphies for considered this family to be of doubtful the family, comprising only three genera. monophyly. Some cyrtaucheniids are the only . The monophyly of Migidae ap- rastelloids to have two rows ofteeth on broad pears well supported (see Raven, 1985; Go- paired tarsal claws (jointly with female palpal loboff and Platnick, 1987; Goloboff, 1991), claw teeth located prolaterally; figs. 8, 9, 13). and no important apomorphies are shared by Some aporoptychine cyrtaucheniids also share only some migids and any non-migid taxon. with some Tuberculotae the presence of a However, some characters (such as the male serrula, scopula, and corrugiform bothria. tibial apophysis, or the presence of an inter- Five cyrtaucheniid taxa were included in the cheliceral tumescence) show variation within analysis: Bolostromus and FuJius (two typical Migidae, and two genera, Calathotarsus and Aporoptychinae), Myrmekiaphila (Eucteni- Heteromigas, are included in the analysis, zinae, a more typical "rastelloid"), Rhytidi- with the purpose of more accurately esti- colus (which Raven included with doubts in mating the homoplasy of those characters. the Aporoptychinae), and Cyrtauchenius Mecicobothriidae. The family is included (Cyrtaucheniinae). as only one terminal; Gertsch and Platnick . Although there are several char- (1979) and Raven (1985) proposed synapo- acters which support the monophyly of this morphies for the group. Eskov and Zonsthein family, some genera share possible synapo- (1990: 352) stated that "the spermathecae of morphies with taxa outside Idiopidae. Neoc- all the mecicobothriids are apomorphously teniza shares several apomorphies with both modified," and that this supports the mono- Migidae and ; some arbani- phyly of the family. However, they actually tines share the presence of a scopula with refer to autapomorphies in the spermathecae cyrtaucheniids; and the Idiopinae share with of each mecicobothriid genus, which might, the and Migidae the presence of at best, support the monophyly ofeach genus strong digging spines on the anterior legs. For (and not the family). this reason, and also because for some place- Microstigmatidae. There are currently two ments of the family the main idiopid aut- subfamilies included in this family, Micros- apomorphy (bulb with second haematodocha tigmatinae (Raven and Platnick, 1981) and extending below embolus) might be better Micromygalinae (Platnick and Forster, 1982). considered as a plesiomorphic retention, three The Microstigmatinae differ from the Mi- taxa (Neocteniza, Idiops, and Misgolas) were cromygalinae by having the superior tarsal included to test the monophyly of Idiopidae. claws broad and bipectinate (and the female Ctenizidae. The monophyly of Ctenizidae palpal claw has teeth located prolaterally) and was also considered doubtful by Raven (1985: by lacking the ALS. These characters are also 57, 66). The only characters thought to sup- found in some non-microstigmatid taxa. Ra- port the monophyly of the group (female su- ven (1985) and Griswold (1985) have sug- perior tarsal claws with only one strong tooth gested that microstigmatids, and especially and anterior legs with strong digging spines) Micromygale (the world's smallest mygalo- might be synapomorphies at a level higher morph), may be neotenic in many characters. than Ctenizidae. However, no characters were Raven (1985) proposed that, plesiomorph- found which would resolve the relationships ically, the Microstigmatidae have the male of the ctenizid groups; no important apo- palpal bulb parallel to the cymbium; that con- morphies seem to be shared by only some dition is approached only in Micromygale. ctenizid and other non-ctenizid taxa. In the Microstigmata has a palpal bulb inserted or- absence ofcharacters, the addition of several thogonally to the cymbium. Although the bulb ctenizid taxa would only decrease resolution; in Pseudonemesia is inserted apically, the bulb therefore, only one genus is included (Um- itself is not directed apically (as in mecico- midia). The results of the present analysis bothriids, atypids, and antrodiaetids), but to- therefore cannot test the monophyly ofCten- ward the base; the tarsus is very long, basally izidae. excavated, and the bulb (smaller than in most Actinopodidae. Actinopus and Plesiolena mygalomorphs, see Raven and Platnick, are included. Raven (1985) and Goloboffand 1981) rests in that cymbial cavity (fig. 3). In 6 AMERICAN MUSEUM NOVITATES NO. 3056 most mygalomorphs the bulb rests instead in of the group. Therefore, four genera are in- a tibial cavity. The cymbial shape and type cluded in the analysis: Scotinoecus, Atrax, ofbulb insertion found in Pseudonemesia are Hexathele, and Porrhothele. thus different from those in any other my- Dipluridae. The only characters supporting galomorph, except perhaps Micromygale and the monophyly of this family are the PLS Ministigmata (and possibly some Anamini; widely separated, with a long apical article, see Raven, 1984: figs. 38, 39). In the latter and the low, hirsute thorax (Raven, 1985: fig. two genera, the cymbium is very short but, 6; the cladogram shows only the latter two as in Pseudonemesia, the apex of the cym- characters). The second character is too bium lacks the evident notch present in most vaguely defined and, because ofdifficulties in mygalomorphs. Some uncertainty exists also scoring, is not considered here (see below). regarding the homology between the meci- Some diplurids (Diplurinae) share with taxa cobothriid conductor and that of Micromy- outside the family several characters; Raven gale (considered as doubtful by Raven, 1985). (1985: 51) discussed the possibility ofa closer Some species ofPseudonemesia (or a closely relationship between diplurines and neme- related genus) have a form of paraembolic siids. Because the monophyly of the family apophysis which is almost certainly not ho- seems debatable, four genera currently as- mologous with the mecicobothriid conduc- signed are included in the analysis: Diplura tor, and the structure which has been consid- (which shares several apomorphies with some ered a "conductor" in Micromygale may well Crassitarsae), Ischnothele, Chilehexops, and represent a modification of that apophysis. Euagrus. A detailed examination ofthe palps ofMi- Nemesiidae. As mentioned above, it is cromygale and Ministigmata was not possi- plausible that the characters used by Raven ble, because of the paucity of available spec- (1985) to support the monophyly ofthe fam- imens and their tiny size, and scoring ily might actually be plesiomorphies. Several Micromygaleforpalpal characters proveddif- nemesiid genera, representing most of the ficult. The bulb and cymbium in Micromy- subfamilies proposed by Raven (1985), are gale and Ministigmata may represent an ad- included in the present analysis: Stenoter- ditional modification of the state found in ommata, Ixamatus, Acanthogonatus, Ne- Pseudonemesia, or they may represent an in- mesia, Xenonemesia (which could not be dependent modification arising from a more placed in any of Raven's subfamilies; Golo- plesiomorphic type of palp. Given the un- boff, 1988), and Neodiplothele. The genus certainties regarding palpal characters in Mi- Neodiplothele was tentatively included by cromygale, at the present time it seemed wis- Raven (1985) in Pycnothelinae; it shares sev- er to code that genus as having a palp eral possible apomorphies with some thera- representing additional modifications from phosoid taxa (the presence of clavate tarsal those found in Pseudonemesia. Such scoring trichobothria, two rows of tarsal trichoboth- was admittedly based not only on actual ria, and the characters mentioned by Raven, observations, but also to some degree on ex- 1985: 103). I examined the types of Neodi- isting hypotheses of relationships, and is plothele irregularis Mello-Leitao and N. flu- therefore arguable. Representatives of Mi- minensis Mello-Leitao only at an early stage crostigmatidae included are the genera Pseu- ofthis study and therefore some of the char- donemesia, Micromygale, and Microstigma- acters could not be scored for that genus. ta, together with some currently undescribed Barychelidae. No unequivocal synapo- genera (see below). morphies for this family were proposed by Hexathelidae. Only one character, the Raven (1985); the characters that he pro- presence of numerous labial cuspules, has posed would be, under some placements of been proposed to support the monophyly of the family, simply plesiomorphies. However, this family (Raven, 1978, 1980, 1985). Be- there seems to be a character which unam- cause some apomorphies (loss of ALS, for biguously supports the monophyly of the example) are shared by some non-hexathel- group: the anterior rim ofthe booklung open- ids and some but not all Hexathelidae, there ing has a series of teeth (forming a kind of may be good reason to doubt the monophyly comb) (figs. 4, 5). This character (absent in 1993 GOLOBOFF: REANALYSIS OF MYGALOMORPHS 7 all the non-barychelid genera I have exam- true ofXenonemesia). These new genera will ined) is present in trichopelmatines (consid- be described shortly (in collaboration with N. ered by Raven, 1985, as the sister group of Platnick); they are included in the data matrix all the other barychelids), sasonines, and bar- as "New genus from Ecuador" and "New ge- ychelines. Raven (in litt.) also has found that nus from Mexico." the tarsal organ of barychelids is placed api- Rooting. The tree was rooted using Me- cally on the tarsus (whereas other mygalo- sothelae. Although the sister group of my- morphs have the tarsal organ at a distance galomorphs is the Araneomorphae, they are from the tip), which appears to be an addi- so highly modified that in only a few char- tional synapomorphy for the group. Bary- acters (detailed below under Characters) could chelidae is therefore included as one terminal they be used for comparison. For the char- in the analysis. The ancestral states for the acters which are comparable in araneo- family were inferred under the assumption morphs but variable within that group, Plat- that trichopelmatines are the sister group of nick et al.'s (1991) and Coddington and Levi's the other barychelids (as proposed by Raven, (1991) cladograms were used to determine 1985), with the exception (for the reasons the plesiomorphic states for araneomorphs. indicated below under Characters) of char- CHARACTERS acter 13. Probably our understanding of the Barychelidae will change substantially with The characters in the following listing are ongoing revisionary studies by Raven. tabulated in the data matrix of table 1. All Theraphosidae. One of the aims of the the multistate characters were considered present analysis is to test whether the pres- nonadditive; in most cases, there is no evi- ence of two rows of teeth is actually a syna- dence for ordering the states, but when some pomorphy for nemesiids. Some male thera- evidence for ordering might exist, the reasons phosids have two rows ofteeth on the superior why the character is considered as nonaddi- tarsal claws. The family is therefore repre- tive are discussed (i.e., characters 13, 21). sented in the present analysis by two taxa, Character 0: Thorax flat = 0; thorax slop- Ischnocolus (for which the males have two ing = 1. rows of teeth), part of a basal unresolved te- Character 1: Eyes sessile = 0; eyes on a trachotomy in Theraphosidae, and Thera- common tubercle = 1. Although the eyes in phosinae (in which both sexes have unarmed are set on a tubercle, the outgroup claws). Theraphosinae is a group ofuncertain is coded as missing (i.e., having either state), limits, due to the large number ofgenera, but because araneomorphs have sessile eyes. the inclusion of many genera in this group is Character 2: Serrula absent = 0; present = clear (see Raven, 1985: 118). 1. Raven (1985) considered that the structure . The family Paratropididae found on the maxillary lobe ofliphistiids was was enlarged by Raven (1985) to include the not homologous to that in mygalomorphs. genus Melloina (= Glabropelma). Melloina However, in both mygalomorphs and liphis- and Paratropidinae are included as two ter- tiids the structure is-although not identi- minals. cal-relatively similar and located in the same Currently unplaced taxa. Two new genera position. The structure in mygalomorphs and (from South America) are included in the Mesothelae may or may not be homologous, analysis. Both have two rows ofteeth on broad and the state in Mesothelae is therefore coded superior tarsal claws, and a female palpal claw as missing. Because the ixamatines include with teeth located prolaterally, which would both serrulate and aserrulate species (Raven, in principle indicate affinities with Nemesi- 1982), the fit for this character was calculated idae. However, placing those genera in the counting one additional step. Nemesiidae seemed as parsimonious as plac- Character 3: Tarsal spines present = 0; ab- ing them in the Microstigmatinae, which also sent = 1. Araneomorphs have state 1, but the have two rows of teeth. The booklung spir- Mesothelae and Amblypygi have state 0; the acles of these genera, although not as modi- outgroup is therefore coded as 0. fied as in Microstigmata, are smaller and more Character 4: Labium short = 0; subquad- rounded than in most nemesiids (the same is rate = 1; long = 2. One step within terminals 8 AMERICAN MUSEUM NOVITATES NO. 3056

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0)

0 0 1993 GOLOBOFF: REANALYSIS OF MYGALOMORPHS 9

counted, because barychelids include species with state 0 and species with state 1. Character 5: Axis of bulb parallel to cym- bial axis = 0; orthogonal or directed toward Co r-I 0 00 00 0 the base = 1. 0 0 00 0 Character 6: Maxillary cuspules absent = -- 0 000 t')" 00 0I)0I)0n000 0; present = 1. The character is variable in iOa 0 0 0 0 00 0 and thus one within 0) atypids, step terminals coOD 00 0 0 0o 0Io 0o 0 was added to this character. 0o Character 7: ALS present = 0; absent = 1. Co Character 8: Thoracic fovea an open pit 0 '-'40 (sometimes longitudinal) = 0; transverse = CD EOo -)I0O)'-4 )00 1; very wide = 2; closed and longitudinal = 0)9 ')O 3. Most mygalomorphs have states 1 or 2. .-A ,-4 0\ *\ 020 fi C)O C' C) 0 The problems in scoring this character are co -I0- Or- O CO'-I C CCl C)2 Cl C'2 mainly in the plesiomorphic groups of my- a:oO galomorphs. Antrodiaetids have state 3; I 0) oO O C) C 00 O O consider mecicobothriids as having state 0 O H C)00O Co) O O O 0 C and diplurids and hexathelids as having O Or- C)'-4O-4 C)O'-4 C>2 C) state 1. O'-4 OC 0) 0 O O C) C Character 9: ITC dentate =0; edentate = 1. 0) Character 10: ITC normal = reduced in isCDEO O C) 0; a CO)00O C1 1- r- _I C) size (or absent) = 1. C00 EIO Character 11: Female tarsi I-II without .6 00 C) O C> C) C ;3 00O 0 C) 0 O C) scopula = 0; scopulate = 1. 00O r- 0 CO 0 z 00O Character 12: Claw tufts absent = 0; pres- O r- O ODOD O00000oC) 0 0 ent = 1. -- isI-EO C CD9 Character 13: Superior tarsal claws. Raven Co 00000o O) O r-I00000HO- -I O4 O- (1985) used the STC to CO OI 00000o dentition of support OD O-I -I O--4 his cladogram. I consider the broadening of 00C0 ~, 4 r- H r 0 '- risd1 00~ the STC as a necessary correlate of the pres- 00~ C) C) O C) ce I~ )t, ence of two rows of teeth, and therefore do 00~ 0 O O O- C) not consider it a separate character. Raven 00~ 0 00 0O 00~ (1985) included as an additional character for 0)OD nemesiids the teeth ofthe palpal claw located HCD prolaterally. The female palpal claw teeth tend, in fact, to be located toward the pro- C-2 C\z C C'O C' C' C- C> r- 0 lateral side of the claw, even in taxa where H C'2 C\ the leg claws have only one row ofteeth (figs. 0 O 00 0 O C0O 10, 1 1). That displacement is even more ev- Cd o o u ident in those taxa in which the leg claws are bipectinate (figs. 8, 9, 12, 13). Thus, I con- C2 sider the displacement of the teeth in the fe- F- male palpal claw and the double row ofteeth as the same character. sH The alternative states for the taxa included in the analysis are: EE f State 0: in Mesothelae (and, plesiomorph- ically, in araneomorphs), antrodiaetids, atyp- ids, non-diplurine diplurids, hexathelids, mecicobothriids, and micromygalines, there is a single row of several teeth; 10 AMERICAN MUSEUM NOVITATES NO. 3056

".

:.,; k ,1j 5 r , II

Figs. 4-7. 4, 5. Strophaeus sp. (Peru), left booklung and booklung comb. 6. Ummidia sp. (Mexico), bothrium from tarsus II. 7. Rhytidicolus sp. (Venezuela), tarsal organ from tarsus I. State 1: in Ischnocolus (Theraphosidae) and is only one strong tooth on the STC (the other many barychelids, males have two rows, and idiopids included in the data set, the migids, females have only one row, of few, minute and Myrmekiaphila, also seem to have this teeth; state, or a homolog of it); State 2: in nemesiids, microstigmatines, di- State 4: in most Theraphosidae (including plurines, and aporoptychines, both sexes have Theraphosinae), some barychelids, and Mel- two rows ofteeth; Raven (in prep.) describes loina there is only one row of a few, minute a new barychelid genus in which both sexes teeth in both sexes. have two rows ofteeth, and suggests that this Males ofRhytidicolus are unknown; the ge- could be the plesiomorphic state for bary- nus is coded as a missing entry (presumably chelids (that suggestion is followed here); males have two rows of teeth on the STC). State 3: in paratropidines, ctenizids, some In Bemmerinae (currently included in Ne- idiopids (Neocteniza), and actinopodids there mesiidae) males have one row of teeth and 1993 GOLOBOFF: REANALYSIS OF MYGALOMORPHS I1I

Figs. 8, 9. Bolostromus sp. (Peru), left female palpal claw. females have two; no bemmerines are in- the present analysis is a stronger test of Rav- cluded in the present data set, and so the state en's cladogram (i.e., an additive coding would is not considered further. imply more steps against Raven's tree). Given the differences between sexes, an ob- Character 14: Maxillary lobe unmodified vious possibility seems to be coding this char- = 0; anteriorly produced = 1. acter separately for males and females. Doing Character 15: Few or no labial cuspules so, however, would imply more instances of = 0; numerous labial cuspules = 1. homoplasy for Raven's (1985) familial clado- Character 16: Caput low =0; elevated = 1. gram. For example, given that nemesiids, di- Character 17: Rastellum absent = 0; pres- plurines, and microstigmatids have bipecti- ent = 1. nate STC in both sexes, such coding would Character 18: Anterior and posterior legs imply that claws provide twice the number of approximately the same size = 0; anterior ofextra steps against Raven's cladogram (i.e., legs shorter and more slender than posterior that the double rows of teeth was acquired legs = 1. independently in males and females). Each Character 19: Spines on posterior legs dis- one ofthe five possible dentitions for the STC tributed ventrally as well as dorsally = 0; lo- was considered as an alternative state of the cated on dorsal surfaces = 1. same, nonadditive (unordered) character, so Character 20: Female tarsi slender = 0; that each state can transform into any other stout = 1. state with equal cost. This coding is not too Character 21: Second haematodocha ex- constraining ofthe possible outcomes; for ex- tending below embolus = 0; not = 1. The ample, if Ischnocolus and barychelids were males of the new microstigmatid genus from to be placed within a group having only one Ecuador have a strongly modified bulb, with strong tooth in the STC (as in ctenizids and a very inflated second haematodocha; be- some idiopids), only one step and no ho- cause this condition may or may not be ho- moplasy would be counted, even when the mologous with that in idiopids (although ap- similarities between male tarsal claws among parently the membranes do not extend below Ischnocolus + barychelids and nemesiids, the embolus as much as in that group), the microstigmatines, etc. are being ignored by state is coded as missing. such a cladogram. Although less informative, Character 22: Apical article of PLS short, the nonadditive coding was preferred, so that domed = 1; digitiform = 0. The character is 12 AMERICAN MUSEUM NOVITATES NO. 3056

2!II},

Figs. 10-13. Female palpal claws. 10. Porrhothele antipodiana, right palp. 11. Atypoides gertschi, left palp. 12. Diplura sp. (Bolivia), right palp. 13. Fufius sp. (Bolivia), right palp.

variable within Acanthogonatus, and so one (because microstigmatids have scales and additional step was counted when calculating pustules and ixamatines only have the latter), the fit for this character. they are coded here as having the same state. Character 23: Cheliceral furrow with two Character 25: Dorsal abdominal tergite rows of teeth = 1; teeth only on promargin present = 0; absent = 1; abdomen with dorsal -0. scutum = 2. Raven (1985) considered the Character 24: Leg cuticle scaly = 1; smooth scutum in some microstigmatids as "ques- - 0. Although Raven and Platnick (1981), tionably homologous" with the abdominal and Platnick and Forster (1982) considered tergites. By the present coding and nonad- that the cuticular modifications in ixamatines ditive interpretation for this character, the were different from those in microstigmatids homology between tergites or scuta, or the 1 993 GOLOBOFF: REANALYSIS OF MYGALOMORPHS 13

Figs. 14-17. Spigots of PLS. 14. Ixamatus broomi. 15. Fufius sp. (Bolivia). 16. Acanthogonatus sp. (Chile). 17. Stenoterommata sp. (Argentina). lack of it, can be decided by congruence with counted forAcanthogonatus, variable for this other characters. character. Character 26: Female anterior tibiae and Character 30: Bothria corrugiform = 1; metatarsi with digging spines = 1; with nor- smooth = 0. Corrugiform bothria appear to mal, elongated spines = 0. be much more widely distributed than pre- Character 27: Ocular quadrangle wide = viously thought; in the present study, they 1; narrow = 0. were observed in Fufius, Bolostromus, Rhy- Character 28: Male palpal bulb with con- tidicolus, Neocteniza, and Ummidia (fig. 6); ductor = 1; without conductor = 0. Idiops has smooth bothria. Character 29: Male tarsi pseudosegmented Character 31: Second haematodocha ex- = 1; integral = 0. One step within terminals tending below embolus = 1; small = 0. 14 AMERICAN MUSEUM NOVITATES NO. 3056

Character 32: Male palpal tibia with thorn- lus, and Bolostromus have a low tarsal organ like spines = 1; with normal spines or un- with concentric ridges; those ridges are al- armed = 0. most inconspicuous in Ummidia, Rhytidi- Character 33: One cymbial lobe pointed = colus, and Idiops. As with character 30, the 1; palpal tarsus normal = 0. state previously thought to define some re- Character 34: Multilobular spermathecae. stricted groups (i.e., state 1; see Raven, 1985: Raven (1985) considered this character to be 22) is actually quite widely distributed. one of the synapomorphies for Cyrtauchen- Character 39: PLS with spigots on medial iidae. By multilobular Raven apparently re- and basal articles only = 0; spigots on the ferred to the presence of multiple divisions three articles of the PLS = 1; spigots only on of each spermathecal receptaculum; this the apical article (if present on the medial eliminates taxa having paired undivided re- article, only on the apical edge) = 2. Palmer ceptacula (what has been called "2 + 2 sper- (1990) reported paratropidids as lacking spig- mathecae"; Platnick and Gertsch, 1976). ots on both the basal and medial article of Several taxa outside Cyrtaucheniidae were the PLS. However, she examined only one scored as having multilobular spermathecae species of Anisaspis; in Paratropis (and also (state 1): Diplura (see Goloboff, 1982b), Is- Melloina) there are numerous spigots on the chnothele, Porrhothele, and Ischnocolus. The basal and medial article (as well as the apical, plesiomorphic state ofthis character for Sten- which is not known to lack spigots in any oterommata is presumed to be multilobular mygalomorph); the plesiomorphic state for (as in S. platense Holmberg, see Goloboff, paratropidines is therefore considered to be 1982a); because some species in the genus basal and medial article with spigots. have undivided receptacula, one step within Character 40: Low parallel ridges on the terminals is counted for this character. male palpal bulb = 1; no ridges = 0. Character 35: Anterior tarsi very densely Character 41: Intercheliceral tumescence scopulate = 1; less densely scopulate = 0. present = 1; absent = 0. The character is Character 36: PLS widely separated = 1; variable in Nemesia and the barychelids; two close = 0. Some species of Hexathele have additional steps were counted to calculate the the spinnerets more separated (see Forster fit. and Wilton, 1968), and therefore one addi- Character 42: Tarsi with clavate tricho- tional step was counted when calculating the bothria = 1; without = 0. Liphistius (Me- fit for this character. sothelae) has clavate trichobothria, but Hep- Character 37: Booklung openings normal tathela, the other genus in the Mesothelae, -0; small and rounded = 1 (Raven and Plat- has only filiform trichobothria; this would nick, 1981: figs. 1, 2). The booklung openings suggest scoring Mesothelae as a missing entry ofthe new genera from Mexico and Ecuador, (i.e., having either state). In Liphistius, how- and those ofXenonemesia, were coded as for ever, there are alternating, distinct rows of Microstigmata, Pseudonemesia, and Mi- clavate and filiform trichobothria (Murphy cromygale despite some apparent differences. and Platnick, 1981; Platnick and Sedgwick, In the three former genera the booklung 1984). In the theraphosoids and some cten- openings are not as small, and the posterior izids and idiopids, in contrast, the clavate margin is not as sclerotized as the anterior trichobothria are interspersed among the fi- one, as is the case for the other three genera. liform ones. The state in Liphistius is thus All of them, however, seem to share a com- considered nonhomologous with that in ther- mon type of modification (which progresses aphosoids and some rastelloids, and because further in Microstigmata and Pseudoneme- clavate trichobothria are also absent in ara- sia, and possibly Micromygale), and are neomorphs and Heptathela, the outgroup is therefore coded as having state 1. coded as zero. Character 38: Tarsal organ low and smooth Character 43: A strongly curved row of - 0; low with concentric ridges = 1; domed trichobothria on the anterior metatarsi, ex- and smooth = 2. The distinction between tending retrolaterally (sometimes the basal states 0 and 1 is sometimes difficult; Neoc- trichobothria separated from the others) = 1; teniza, Idiops, Ummidia, Fufius, Rhytidico- a straight or gently curved dorsal row = 0. 1 993 GOLOBOFF: REANALYSIS OF MYGALOMORPHS 15

Character 44: Tarsal trichobothria in a zig- Character 48: Posterior lateral spinnerets zag row = 0; forming a wide (usually diago- long = 1; short = 0. Raven (1985) suggested nal) band = 1; in two longitudinal lines sep- that the spinneret elongation in diplurids was arated by setae = 2; reduced = 3; in a straight of a different nature than that in hexathelids row = 4. In theraphosids, non-paratropidine and mecicobothriids, which would suggest paratropidids, barychelids, and Neodiplothe- scoring them differently. According to Ra- le the tarsal trichobothria form two zig-zag ven, in diplurids the three articles ofthe PLS rows, separated by a band of setae. The tri- are elongated, whereas in the other two groups chobothria of both the theraphosoids and only the apical article is elongated. The dis- several rastelloid taxa have been previously tinction between the two types, however, is described as a "broad band oftrichobothria" not clear for some of the species included in (see Raven 1985: 116); in rastelloids, how- the present data set (e.g., the relative lengths ever, there is no medial row of setae, and of the PLS segments in Scotinoecus and Eu- those taxa (Actinopus, migids, ctenizids, agrus are essentially the same), and the me- Idiops, Neocteniza, and Myrmekiaphila) are cicobothriids and long-spinnereted hexath- here considered as having a different state. elids are therefore scored as having the same The tarsal trichobothria ofparatropidines do state as the diplurids. not fit exactly in either states 1 or 2. Although Character 49: Spigot types (Palmer, 1990): there is no medial row of setae, the tricho- articulated = 0; pumpkiniform = 1; fused = bothria form a longitudinal band. The state 2. Among the taxa included in the data set, in paratropidines is here considered as a fur- those studied by Palmer (1990), except Sten- ther modification of the state in the other oterommata andAcanthogonatus, were scored theraphosoids (i.e., state 2); because that on the basis of her study. For the present unique state would be uninformative within study, the spinnerets of the following taxa theraphosoidina, paratropidines are scored were examined (figs. 14-19): six species of as having state 2. Acanthogonatus [A. fuegiana (Simon), the Raven (1985: 143) erroneously described species identified as A. frankii Karsch by the family Actinopodidae as having "a more Palmer, 1990, plus four undescribed species], or less straight row" of trichobothria on the three undescribed species of Stenoterom- tarsi. That statement is true only ofPlesiolena mata, Ixamatus broomi Hogg, Nemesia du- and Missulena; Actinopus has a wide band of bia 0. P.-Cambridge, Xenonemesia platensis trichobothria (Goloboff and Platnick, 1987). Goloboff, and unidentified species of Di- Character 45: Palpal coxae subquadrate = plura, Rhytidicolus, Fufius, and Bolostromus, 1; elongate = 0. plus a variety of"nemesiids" not included in Character 46: Spermathecae paired (2 + the data set [Pycnothele singularis (Mello- 2) = 0; unpaired = 1; nonreceptaculate = 2. Leitao), Lycinus epipiptusa (Zapfe), Diplothe- Although some species of Stenoterommata lopsis bonariensis Mello-Leitao, an unde- have 2 + 2 spermathecae (see Goloboff, scribed genus of Diplothelopsini, Chaco 1982a), other species in that genus have un- obscura Tullgren, and two undescribed spe- paired spermathecae, and I consider the latter cies ofChaco]. As discussed by Palmer (1990), to be the plesiomorphic state for the genus. classifying a particular spigot as being One step within terminals is counted. "pumpkiniform" or "articulate" is some- Character 47: More than three spines on times equivocal. According to Palmer (1990), patella III = 1; 3 or fewer = 0. Ctenizids, Acanthogonatus lacks pumpkiniforms, but all actinopodids, idiopids, and cyrtaucheniids the species ofAcanthogonatus examined for have numerous prolaterodorsal spines on pa- this study (including specimens collected in tella III. Those taxa also have the spines on the same locality as Palmer's, and probably the posterior legs situated dorsally, a possibly conspecific) have what I consider to be clear associated character. In some species, how- examples of pumpkiniform spigots. Those ever, one of the characters is present and the pumpkiniforms are larger than the articulat- other is absent, and the presence of dorsal ed spigots, and are all located on the inner spines and the number of patellar spines are edge of the spinning field (figs. 16, 18) (see therefore considered independent characters. character 65). 16 AMERICAN MUSEUM NOVITATES NO. 3056

Character 50: Spigot shaft sculpture (Palm- debate on several aspects of the spider post- er, 1990): overlapping scalelike folds = 0; embryonic development (including the no- minimal surface detail = 1; pointed projec- menclature of the stages). I will follow Ga- tions = 2. The taxa examined for character liano (1969, 1972, 1973a, 1973b) in 49 were also examined for this character. considering that the first postembryonic in- Character 51: Slit on spigots present = 0; star may occur before hatching (i.e., before absent = 1. Palmer (1990) mentioned this the chorionic membranes are broken). Ga- character but did not include it in her table. liano (personal commun.) has kindly provid- The character was scored based on her pho- ed a summary of the published studies and tographs. unpublished observations she has made on Character 52: Cheliceral fangs short, thick, the development of Actinopus insignis and diagonal = 1; long and parallel = 0. (Holmberg), Diplura paraguayensis (Schia- Character 53: Sternum gradually narrowed pelli and Gerschman), Oligoxistre argenti- in front = 0; sternal sides more parallel = 1. nensis (Mello-Leitao), Xenonemesia platen- In the Ctenizidae, Actinopodidae, Idiopidae, sis, and an undescribed Stenoterommata. My Migidae, and Antrodiaetidae, the sternum scoring for this character, as well as that for narrows gradually from coxae III; the labium character 60, was based on Galiano's sum- is thus approximately as wide as the anterior mary. I have assumed that the two plesiom- end of the sternum. In the other mygalo- orphic theraphosids Oligoxistre (placed by morphs, except Atypidae, the sternum is ap- Raven in the possibly paraphyletic Ischno- proximately as wide between coxae II as it is colinae) and Ischnocolus share the same type between coxae III; the labium thus appears of postembryonic development. According much narrower than the sternum. The Me- to Galiano, all mygalomorphs except Gram- sothelae and Atypidae have sternal shapes mostola have three postembryonic instars [the that are very different from those, and among presence offour instars in Grammostola (Ga- themselves; because both of those unique liano, 1969, 1973a) is easily interpretable as states would be uninformative, those taxa are an autapomorphy]. There is variation, how- scored as missing entries. ever, in the degree of development of the Character 54: Anterior legs with scopula stages. Ischnothele, Diplura, Atypus, and An- symmetrical = 1; scopula more developed on trodiaetus (and Heptathela) have a second the prolateral side = 0. In barychelids, scop- postembryonic instar more developed, with ulate idiopids, and some scopulate cyrtauch- cephalothorax and abdomen in the same eniids, the scopula on the anterior tarsi and plane, whereas Actinopus, theraphosids, and metatarsi is more developed on the prolateral "nemesiids" have the cephalothorax and ab- side. In the Diplurinae, Nemesiidae, Apo- domen at 900 (only later instars have the roptychinae, and Theraphosidae, in contrast, cephalothorax and abdomen in the same the scopula is equally developed on both sides. plane). The nonscopulate taxa are scored as missing. Character 60: Cheliceral fang with apical Character 55: Postlabial sigillum deeply tooth in larval stages = 0; with simple, conical excavated = 1; a shallow suture = 0. claw in all stages = 1. Diplura and Ischnothele Character 56: Spermathecal ducts more have a simple claw; all the other studied my- strongly sclerotized basally = 1; with uniform galomorphs have a toothed fang (see Galiano, sclerotization = 0. 1973b: 325 for comments on Antrodiaetus). Character 57: Fovea sinuous, recurved = The outgroup is coded as 1 because Heptathe- 1; straight or procurved = 0. la has a simple claw in the chelicerae (Yoshi- Character 58: Spinnerets well separated kura, 1955, quoted in Galiano, 1973b). from anal tubercle = 0; spinnerets and anal Character 61: Cymbium with apical rim tubercle close = 1. sclerotized (bulb cavity apically closed) = 0 Character 59: Second postembryonic in- (fig. 1); cymbium apically incised and mem- star well developed, with cephalothorax and branous (bulb cavity apically open), enclos- abdomen in the same plane = 0; less devel- ing subtegulum = 1 (fig. 2); cymbium with oped, with cephalothorax and abdomen per- apical edge membranous, but not incised and pendicular = 1. There is currently an ongoing not enclosing subtegulum = 2 (fig. 3). See 1993 GOLOBOFF: REANALYSIS OF MYGALOMORPHS 17 above, Taxa, for discussion ofthe coding for (figs. 20, 21). This structure is difficult to ob- Micromygale and Pseudonemesia. serve in many cases; it appears to have a Character 62: Maxillae normal = 0; max- scattered distribution (it was observed in bar- illae concave in the middle (as in paratropi- ychelids, Neocteniza, and Antrodiaetus; par- dids) = 1. atropidines seem to have a modified opening; Character 63: Patella III without apical it appears to be absent in theraphosids, ne- comb ofspines = 0; with apical comb ofspines mesiids, diplurids, and Idiops). Given that (Platnick and Shadab, 1976) = 1. the distribution of this character in mygalo- Character 64: Bothria normal = 0; with a morphs is still poorly known, it is not in- sinuous impression around tricheme aper- cluded in the analysis. ture (Goloboff and Platnick, 1987) = 1. Another possibly informative character is Character 65: Pumpkiniform spigots dis- the presence of a peculiar type of cuticular persed (or absent) = 0; forming a row in the modification near the maxillary glands of inner edge of spinning field = 1. State 1 is some genera (figs. 22-25). The basal part of shared by Acanthogonatus (figs. 16, 18) and the maxillae (the part in contact with the la- Stenoterommata (fig. 17; Palmer, 1990, er- bium) has a less sclerotized, paler, and gen- roneously stated that the pumpkiniforms in erally well defined area (in many cases pos- Stenoterommata are in a "medial row;" the teriorly delimited by fringes of modified row is actually lateral, along the inner edge hairs). In that paler area most of the exam- of the spinning field). ined genera have conical hairs, with a blunt Character 66: Labium normal = 0; labium tip; a glandular secretion apparently would big, squared, and very inclined = 1. exit from an opening in the base of the hair; Character 67: Male tibial apophysis: no the hairs are only visible with scanning elec- apophysis = 0; with apical prolateral mega- tron microscopy. Those hairs have been ob- spines (as in Hexathele and Scotinoecus) = served in Antrodiaetus unicolor (Hentz), Me- 1; on leg II = 2; a retrolateral apical mega- cicobothrium thorelli Holmberg, Hexathele spine = 3; theraphosoid type of tibial spur hochstteterri (Ausserer), Stenoterommata (Raven, 1985) = 4; idiopid type oftibial spur platense, and unidentified species of Stro- = 5. phaeus (Barychelidae, Barychelinae), Bolos- Character 68: Posterior leg spines normal tromus, Ischnothele (fig. 25), and Diplura (fig. = 0; reduced to spiniform setae = 1. 24). They are absent in Cyriocosmus sp. Character 69: Posterior sternal sigilla nor- (Theraphosidae, Theraphosinae) and in the mal = 0; reduced = 1. two Domiothelina examined, Idiops clarus Character 70: Cheliceral fang normal = 0; (Mello-Leitao) (figs. 22, 23) and an uniden- keeled = 1. tifiedActinopus. The loss ofthe modified hairs may therefore be a synapomorphy of the CHARACTERS NOT CONSIDERED Domiothelina, or some group within Dom- IN THE ANALYSIS iothelina, with parallelisms in the Thera- Raven (1985: 5 1) considered the "lowered phosidae. Since the distribution of the char- caput plus elevated thoracic region" as a syn- acter in mygalomorphs is poorly known, it is apomorphy for diplurids, shared with some not included in the data set. microstigmatids and possibly some bary- The following characters (or scorings of chelids. I have been unable to detect any sig- taxa) were mentioned by Eskov and Zon- nificant difference between the thorax of di- shtein (1990) in support oftheir scheme. For plurids and that ofmost other mygalomorphs. the reasons discussed in each case, those In Pseudonemesia, it is clear that the thorax characters are not included in the data set. is more elevated behind the fovea, as noted Eyes on a common tubercle in atypids. Es- and illustrated by Raven and Platnick (1981), kov and Zonshtein claimed (p. 344) that but for the present choice of taxa Pseudo- atypid eyes are set on a tubercle. Unlike the nemesia would be autapomorphic for this eye tubercle of most of the taxa included by character. Raven (1985) in his Tuberculotae, however, One potentially informative character is the the atypid eye tubercle comprises only the presence of a clypeal opening in some taxa AME and slopes more gradually; atypids are 18 AMERICAN MUSEUM NOVITATES NO. 3056

Figs. 18-21. 18. Acanthogonatus sp. (Argentina), spigots of PLS. 19. Diplura sp. (Bolivia), spigots of PLS. 20, 21. Antrodiaetus unicolor, clypeal opening. therefore scored as not having an eye tubercle ference lies, and I have been unable to detect (contra Eskov and Zonshtein, 1990). any. Therefore, instead of scoring this char- Maxillary cuspules of atypids. Eskov and acter as absent in atypids (as they would have Zonshtein (p. 348) proposed that the maxil- it), the character is scored as either absent or lary cuspules of atypids are different from present (because Calommata, the sister group those in non-atypoid taxa (all mygalomorphs of cuspulate atypids, lacks cuspules and except Mecicobothriidae, Antrodiaetidae, and therefore the ancestral state for this character Atypidae). They claimed that the cuspules in in atypids is ambiguous). atypids are "clearly distinguishable by their Shape of the chelicerae. According to Es- shape from the [non-atypoid] ones." They do kov and Zonshtein (p. 348) the chelicerae in not mention, however, where the actual dif- Atypidae, Antrodiaetidae, and Mecicoboth- 1 993 GOLOBOFF: REANALYSIS OF MYGALOMORPHS 19

Figs. 22-25. Maxillary glands. 22, 23. Idiops clarus. 24. Diplura sp. (Bolivia). 25. Ischnothele sp. (Venezuela). riidae are "strongly gibbose dorsally with a ambiguous as to render it completely unin- well extended basal constriction." Although formative; some kind of pleurital expansion the atypid chelicerae fit this description, the exists in most, if not all, mygalomorphs, and chelicerae of mecicobothriids and antrodi- intermediate degrees of development make aetids seem in fact more similar to those of it too difficult to score. non-atypoid taxa. Because this cheliceral Thoracic fovea. Eskov and Zonshtein character, unique for atypids, would be un- claimed (p. 35 1) that the shape ofthe thoracic informative, it is not included. fovea supports the monophyly of both the Pleurital expansions. This character had atypoid and non-atypoid lineages. They, been found by Gertsch and Platnick (1979). however, contradict themselves when they The scoring of this character would be so say that "a few genera (Microhexura in ther- 20 AMERICAN MUSEUM NOVITATES NO. 3056 aphosoids, Atypus, Sphodros and Aliatypus Raven (1985) is due to a different treatment in atypoids) independently retain the ances- ofthe data. This section describes results ob- tral modality of this structure." tained under equal, unitary weights for all "Structure ofthe male palp." According to characters, which are therefore more com- Eskov and Zonshtein, the structure ofthe male parable with Raven's results. palp supports the monophyly ofboth the aty- Hennig86 produced 82 equally parsimo- poid and non-atypoid lineages. Those con- nious trees, 227 steps long. Neither Tuber- clusions, however, are based on a highly spec- culotae nor Fornicephale, Hexathelidae, Di- ulative reconstruction ofthe evolution ofthe pluridae, Nemesiidae, or Cyrtaucheniidae bulb sclerites. They claim to have identified appeared as monophyletic. Except for the re- homologies between the bulbs in Mesothelae lationships of hexathelids and non-diplurine and mygalomorphs, despite hypothetical ex- diplurids, the trees were quite similar to those changes in position between the sclerites. produced in the final analyses (shown in fig. Since one of the main criteria for homology 26). The strict consensus showed Atypidae is relative position, the identification of the plus Antrodiatidae, Euagrus plus Chilehex- sclerites as "the same" must thus be based ops, Ischnothele, Porrhothele, Atrax, and on the other main criterion: intrinsic simi- Scotinoecus plus Hexathele as successive sis- larity (see Coddington, 1989, and Lipscomb, ter groups ofDiplura plus the rest ofthe my- 1992, for a review of the criteria for homol- galomorphs. Those trees thus require a regain ogy). Eskov and Zonshtein, however, do not of ALS for Scotinoecus plus Hexathele. Di- mention what structural details led them to plura clustered with clade 66 (offig. 26), based recognize the sclerites even after the changes on the shared absence of tarsal spines, two in position they postulated, and in studying rows of STC teeth, scopula, and male tibial the relevant taxa I was unable to find simi- apophysis (those characters were reversed or larities which could support Eskov and Zon- modified in some Rastelloidina, clade 65 in shtein's claims. fig. 26). The shortest trees in which Diplur- Strongly enlarged cymbium. Although in idae was monophyletic required three more both atypids and mecicobothriids the cym- steps than the shortest trees; the shortest trees bium is elongated, the structure in each fam- in which Hexathelidae was monophyletic re- ily is different. In mecicobothriids, unlike quired only one. Making Nemesiidae mono- atypids, the palpal tarsus is long and narrow. phyletic required five additional steps. The The character is considered (contra Eskov and character considered by Raven (1985) as a Zonshtein) not to provide evidence ofmono- synapomorphy of Nemesiidae, the presence phyly for mecicobothriids plus atypids. oftwo rows ofteeth on the paired tarsal claws, Number of cardiac ostia. The number of was a synapomorphy at a much higher level cardiac ostia is known for only a couple of even if Nemesiidae was held monophyletic. the taxa included in the present data set. Un- The microstigmatids appeared as a group til more taxa are examined for this character, within Nemesiidae, as suspected by Raven it is unlikely to provide useful information. (1985: 65). Although the shortest trees in which Di- pluridae, Hexathelidae, or Nemesiidae were RESULTS AND DISCUSSION monophyletic were only slightly inferior to the most parsimonious trees, the case was PRELIMINARY ANALYSES different for other groups proposed by Raven Initial analyses were conducted using (1 985) that did not appear on the cladograms. Hennig86 to find shortest trees with all the Constraining Cyrtaucheniidae to be mono- characters equally weighted. Raven (1985) phyletic required nine additional steps (many defended his cladogram on the basis ofcom- apomorphies shared by Myrmekiaphila and parisons ofsimple step differences; he did not the other rastelloids are absent in aporopty- make explicit reference to weighting. The fi- chines). nal results presented in this paper include Raven's cladogram required 37 additional character weighting, so the question might be steps over the shortest tree found (the taxa raised whether the difference in results from not considered by him, Xenonemesia and the 1993 GOLOBOFF: REANALYSIS OF MYGALOMORPHS 21 new genera from Mexico and Ecuador, were essentially paralleled if all the characters not in their most parsimonious placements). considered by Raven (1985) were deactivat- How much did it cost to make monophy- ed. That resulted in multiple equally parsi- letic both Fornicephalae and Tuberculotae, monious trees, the consensus of which was the two main branches in Raven's clado- consistent with the tree in figure 26, and for gram? Constraining those two groups to be which the non-atypoids did form a mono- monophyletic required 1 1 additional steps. phyletic group; neither Dipluridae, nor Ne- The consensus of the resulting trees would mesiidae, nor Hexathelidae appeared as differ from the one in figure 26 in moving the monophyletic. Constraining all the families Rastelloidina to the basal part of the clado- to be monophyletic, given Raven's charac- gram (with Rhytidicolus clustering with Bo- ters, required eight steps beyond those re- lostromus plus Fufius instead ofbeing the sis- quired for the unconstrained data; constrain- ter group of the other Rastelloidina), and in ing Tuberculotae and Fornicephalae to be having less resolution. The 11 additional steps monophyletic cost four steps, but if all the necessary to make Fornicephalae and Tub- families were constrained to be monophylet- erculotae monophyletic indicate that, on a ic, the shortest trees in which Fornicephalae global analysis under the assumption ofequal and Tuberculotae were monophyletic were weights, those two groups could be clearly only three steps longer than those in which rejected. It is interesting that if the mono- they were not. phyly of all the families were assumed prior Two things can be concluded at this point. to the analysis, the shortest trees would be First, the differences between Raven's clado- only six steps longer than those in which Tub- gram and the shortest trees found were caused erculotae and Fornicephalae were also mono- not so much by the addition ofthe new char- phyletic. If one relied on the monophyly of acters, but by making the analysis global, all the families-as Raven (1985) had to- without constraining the families to be the rejection of Tuberculotae and Fornice- monophyletic. Second, given his data and the phalae would have been much less evident constraints necessary for a hand analysis, Ra- than in a global analysis. ven (1985) was impressively close to an op- The groups Tuberculotae and Fornice- timal solution. phalae were in conflict with the dichotomy of atypoid vs. non-atypoid mygalomorphs. Platnick (1977) had shown that all the char- FINAL RESULTS acters shared by the Atypoidea (sensu Simon, The final analyses included a more careful 1892, i.e., atypids, antrodiaetids, and meci- consideration of the reliability of the char- cobothriids) were plesiomorphies. Because acters (as discussed above under Methods), there were several differences among aty- producing a more realistic treatment. At this poids and non-atypoids, the implication was stage of the analysis, Pee-Wee was used to that the non-atypoids-which had all the search for trees. apomorphic states for the characters differing The data set in table 1 produced 36 trees in both groups-were probably monophylet- ofmaximum fit, 366.9, and 228 steps. Figure ic. Contrary to what might have been ex- 26 shows the strict consensus of those trees. pected after Platnick's study, Raven's (1985) For any ofthe resolutions ofthe trichotomies analysis considered not only the atypoids, but of clades 72 and 52, the characters could be also the non-atypoids, to be nonmonophy- optimized such that there is change along the letic groups. Although Raven ofcourse based branch, but alternative optimizations not in- his rejection of the non-atypoids on several volving change exist, so that no resolution is (previously ignored) characters, his familial actually supported. The polytomy ofclade 64 cladogram (Raven, 1985: fig. 1) shows nu- is the consensus of four parsimonious pos- merous parallelisms between the Fornice- sibilities (shown in fig. 27); each ofthose pos- phalae and Tuberculotae-corresponding to sibilities, again, implies the same number of the characters potentially defining the non- steps for every character in the data set, and atypoids. is potentially supported only by ambiguous The results for the complete data set were optimizations. The fit and weight (i.e., the 22 AMERICAN MUSEUM NOVITATES NO. 3056

ATYPIDAE 76 - Antrodiateus+Atypoides - 174J-: Aliatypus MECICOBOTHRIIDAE '75 - Atrax Porrhothele -69JE:9-67JEHexathele 71JE Scotinoecus L73l: - Chilehexops F72- r Euagrus Ischnothele =70J Diplura IL68jI Plesiolena 44JE Actinopus .=47 = Heteromigas =43JE- Calathotarsus 52- Misbolas r r45 j Neocteniza -48J Idiops - Ummidia r 55JL- Myrmekiaphila L58J- Cyrtauchenius 63 L Bolostromus 5 Fufius Rhytidicolus Nemesia r- Stenoterommata 620E Acanthogonatus K Neodiplothele 60 BARYCHELIDAE l =56 r- PARATROPIDINAE 49-4E Melloina =53l E: Ischnocolus L50O THERAPHOSINAE r-Ixamatus 61jj= Xenonemesia L57J r n.gen.,Mexico L54 =51rfr- n-gen.EcuadorMicrostigmata L46 Micromygale 42J-- Pseudonemesia Fig. 26. Cladogram for taxa in table 1. The four possible resolutions ofthe polytomy ofclade 64 are shown in figure 27. cost ofadding a step under the fitting function ed characters (reported in the preceding sec- used) for each character are shown in table 2. tion); a comparison ofthose trees may clarify Note that the fittest trees are one step lon- the way in which the weighting method used ger than the shortest trees for equally weight- here works. The fittest trees save one step in

e= THERAPHOSOIDINA Nemesia _ 60 Neodiplothele L=64. OE: THERAPHOSOIDINA ?-L E Stenoterommata Neodiplothele =64L T62 Acanthogonatus Stenoterommata Nemesia LFIxam62as+{60 Acanthogonatus ? Ixamatus+MICROSTIGMATIDAE Ixamatus+MIRCROSTITIADAE

E= Acanthogonatus rIxamatus+MICROSTIGMATIDAE I =62 Stenoterommata =64 r-Nemesia =64 60E: THERAPHOSOIDINA ? emE: THERAPHOSOIDINA Neodiplothele {6056 Neodiplothele =[?- r-Nemesia l-?: rStenoterommata _ ?CIxamatus+MICROSTIGMATIEMAE -62F Acanthogonatus Fig. 27. The four resolutions of clade 64; components not occurring in the four trees are indicated by "?." 1 993 GOLOBOFF: REANALYSIS OF MYGALOMORPHS 23

TABLE 2 TABLE 2-(Continued) Character Fits and Weights Step ("Steps" does not include the steps within ter- Char Fit Steps cost minals; "step cost" is the difference in fit produced by adding a step to the character) 48 5.0 2 1.7 49 3.3 6 0.4 Step 50 2.5 8 0.3 Char Fit Steps cost 51 5.0 2 1.7 0 5.0 2 1.7 52 3.3 3 0.8 1 5.0 2 1.7 53 5.0 2 1.7 2 1.4 6 0.1 54 5.0 2 1.7 3 1.3 8 0.2 55 3.3 3 0.8 4 3.3 5 0.4 56 10.0 1 5.0 5 10.0 1 5.0 57 2.5 4 0.5 6 2.0 4 0.3 58 10.0 5.0 7 3.3 3 0.8 59 5.0 2 1.7 8 10.0 3 2.5 60 5.0 2 1.7 9 1.4 6 0.1 61 10.0 2 3.3 10 5.0 2 1.7 62 10.0 5.0 11 2.0 5 0.3 63 5.0 2 1.7 12 5.0 2 1.7 64 10.0 5.0 13 5.7 7 0.7 65 10.0 5.0 14 3.3 3 0.8 66 10.0 5.0 15 5.0 2 1.7 67 4.2 121 0.4 16 3.3 3 0.8 68 10.0 5.0 17 2.5 4 0.5 69 5.0 2 1.7 18 5.0 2 1.7 70 10.0 5.0 19 3.3 3 0.8 20 5.0 2 1.7 21 5.0 2 1.7 22 1.7 S 0.3 characters 15 (numerous labial cuspules), 30 23 2.0 S 0.3 (type of trichobothrial bases), and 36 (sepa- 24 3.3 3 0.8 ration of PMS and PLS), while requiring two 25 10.0 2 3.3 additional extra steps in character 9 (teeth of 26 3.3 3 0.8 27 5.0 2 1.7 the ITC), and one in characters 6 (maxillary 28 10.0 1 5.0 cuspules) and 44 (tarsal trichobothrial pat- 29 1.3 7 0.2 tern). All ofthe characters in which steps could 30 1.7 6 0.3 be saved (as well as character 30) have three 31 10.0 5.0 or more extra steps in both the fittest and the 32 5.0 2 1.7 shortest trees. In contrast, character 15 has 33 10.0 5.0 only one extra step on the fittest tree, and 34 1.7 5 0.3 character 36 only two. The fittest tree, al- 35 10.0 5.0 though slightly longer, saves steps for those 36 2.5 3 0.5 characters which have less homoplasy, and 37 10.0 1 5.0 is therefore to be preferred. 38 3.3 6 0.4 Most of the groupings that were not sup- 39 6.7 3 1.7 ported in the preceding analysis, are not sup- 40 3.3 3 0.8 41 1.7 4 0.3 ported, either, when the reliability of the 42 3.3 3 0.8 characters is taken into account. These groups 43 5.0 2 1.7 are, again, Nemesiidae, Dipluridae, and Cyr- 44 4.4 9 0.4 taucheniidae among the families, and For- 45 3.3 3 0.8 nicephalae and Tuberculotae among the 46 4.0 4 0.7 higher taxa; Microstigmatidae is placed with- 47 2.5 4 0.5 in the paraphyletic Nemesiidae. Hexatheli- dae appears as monophyletic, and the pres- 24 AMERICAN MUSEUM NOVITATES NO. 3056

TABLE 3 Comparison with Other Trees (For all the characters indicated, the trees in which the corresponding group is monophyletic require either one more or one less step than the fittest tree; the difference in fit for those characters is shown in parentheses) Monophyletic Total Characters with taxon fit diff. Characters with worse fit better fit Dipluridae 1.6 3 (0.2); 11 (0.3); 13 (0.7); 44 (0.4); 46 (0.7); 34 (0.3); 36 (0.8) 67 (0.4) Nemesiidae 0.5 12 (1.7); 42 (0.8); 44 (0.4); 50 (0.3) 4 (0.7); 40 (1.7); 41 (0.3) Cyrtauch. 9.4 0 (1.7); 1 (1.7); 13 (0.7); 16 (0.8); 18 (1.7); 45 (1.7); 52 (1.7) 20 (1.7); 23 (0.3); 44 (0.4); 53 (1.7); 54 (1.7); 67 (0.4) Forniceph. and 25.6 3 (0.2); 4 (0.4); 5 (5.0); 6 (0.3); 7 (0.8); 11 (0.3); 2 (0.3); 17 (0.8); 29 (0.1); Tubercul. 13 (0.7); 21 (1.7); 25 (3.3); 28 (5.0); 30 (0.3); 45 (1.7); 52 (1.7) 34 (0.3); 38 (0.4); 39 (1.7); 44 (0.4); 46 (0.7); 49 (0.4); 58 (5.0); 61 (3.3) ence of ALS in Hexathele and Scotinoecus is Dipluridae monophyletic. Those trees ac- a plesiomorphic retention. count more poorly for the absence of tarsal Table 3 shows the difference in fit and steps spines, presence of scopula, double row of between the best tree and trees in which some teeth on the paired claws, zig-zag trichoboth- groups were constrained to be monophyletic. rial pattern, unpaired spermathecae, and api- For some ofthe unsupported groups, a rel- cal retrolateral male tibial apophysis, shared atively small difference in total fit is required between Diplura, Theraphosoidina + Ne- to make them monophyletic. If Nemesiidae mesiidae (clade 64), and (plesiomorphically) is made a monophyletic group (but still al- the Rastelloidina. It should be pointed out lowing microstigmatids to be part ofthe fam- that the advancement of the female palpal ily), the fittest trees account better for the claw teeth, and the widening of the paired labium length, male bulb ridges, and inter- tarsal claws, which were not included in the cheliceral tumescence, but not as well for the data set (on the grounds that they are nec- presence of claw tufts, clavate trichobothria, essary correlates of the presence of two rows and tarsal trichobothrial pattern, in Neodi- of teeth; see above under Characters), would plothele; the former three characters have only strengthen the difference in fit between more homoplasy than the latter three, and so the best trees and those in which Dipluridae the total fit favors the tree in which Neodi- is monophyletic. plothele clusters with Theraphosoidina. The The decrease in fit is even greater to make trees in which nemesiids form a monophy- the Cyrtaucheniidae monophyletic. Myrme- letic group, and the trees in which they do kiaphila shares with the Actinopodidae, not, account equally well for the dentition of Idiopidae, and Ummidia the reduction in the the paired claws, which is plesiomorphic for tarsal claw teeth, the tarsal trichobothrial pat- nemesiids in both cases. Although microstig- tern, and the presence oftwo cheliceral tooth matids are part of Nemesiidae, the presence rows; and Cyrtauchenius shares with that of scaly cuticle (character 24) in microstig- group the shortening of the apical article of matids and Ixamatus is still more parsimo- the PLS and the strengthening ofthe posterior niously optimized as a parallelism, thereby legs. supporting Raven and Platnick's (1981) hy- The characters that traditionally had been pothesis that the cuticular modifications in used in the dichotomy atypoids/non-aty- both groups are independent. poids fit the best trees very well; those char- A greater decrease in fit is required to make acter are disregarded when making mono- 1 993 GOLOBOFF: REANALYSIS OF MYGALOMORPHS 25 phyletic Fomicephalae and Tuberculotae, ence of paired spermathecae (character 46) resulting in a difference of total fit of 25.6. might be plesiomorphic for mygalomorphs Then, according to the present results, the plus araneomorphs. According to the present traditional division Atypoidea/non-Atypo- results, the paired spermathecae are indeed idea suffered only from the problem of par- the plesiomorphic state for mygalomorphs aphyly ofAtypoidea (hardly surprising, since (including non-diplurine diplurids and some that group was based on plesiomorphies), but hexathelids). The change to unpaired sper- not from the problem ofpolyphyly (since non- mathecae defines a large group of mygalo- Atypoidea was based on real homologies). morphs, including diplurines, Rastelloidina, Are those comments to be taken as an im- and Crassitarsae (theraphosids + paratropi- primatur on the views of Eskov and Zon- dids + barychelids + nemesiids + microstig- shtein (1990)? Certainly not. Those authors matids), with a parallelism in some hexath- have recently attacked Raven (1985) and his elids. methodology. They propose that Atypidae, Another character which shows little ho- Antrodiaetidae, and Mecicobothriidae form moplasy, and received a high weight in the a monophyletic group. Although (as shown analysis, is the presence ofspigots on the bas- here) there are good grounds to think that the al article ofthe PLS (character 39). Both Me- rest of mygalomorphs form a monophyletic sothelae and Araneomorphae lack such spig- group, no real evidence for the monophyly ots, as do the Atypidae, Antrodiaetidae, and of Atypidae, Antrodiaetidae and Mecico- Mecicobothriidae; the presence of spigots on bothriidae has been produced by Eskov and the basal article of the PLS is thus one of the Zonshtein (some of the characters they pro- synapomorphies of the non-Atypoidea. Sec- posed are discussed above). In fact, if all the ondary losses of the spigots on the basal ar- families are scored for the characters they ticle of the PLS characterize a group within proposed, the shortest trees do not have that Microstigmatidae (where the spigots of the group as monophyletic (the consensus of the medial article of the PLS are also lost) and 51 equally parsimonious trees includes a the Migidae. This character may shed some monophyletic non-Atypoidea, with Micro- light on the placement of the recently de- mygalinae, Atypidae plus Mecicobothriidae, scribed Triassic genus Rosamygale Selden and and Antrodiaetidae, as their successive sister Gall (1992). Those authors placed the genus groups). The few characters that Eskov and with doubts in the Hexathelidae; the labial Zonshtein themselves propose to use argue cuspules apparently were not preserved, but against their own conclusions. Many char- the spigots were. Selden and Gall stated that acters proposed by Raven (1985) were dis- the spigots "occur ... distally and laterally regarded as irrelevant or simply not men- along the length ofthe PLS" (Selden and Gall, tioned by Eskov and Zonshtein. This 1992: 228). If that means that the spigots obscures, more than it enlightens, the nature were absent from the basal article ofthe PLS, ofthe problem. Although Raven (1985) may the genus should be excluded from the Hex- have failed to find an optimal solution for athelidae (and possibly from the Orthopal- the problem at hand, he certainly tried to pae, as redefined below). All the hexathelids consider all the evidence-and in so doing examined for this study (the genera included greatly facilitated subsequent work on the in the matrix, plus Mediothele, Teranodes, problem. , and Bymainiella) have spigots on the basal article of the PLS. SYNAPOMORPHIES The presence of two rows of teeth (char- acter 13), scopula (character 11), a zig-zag The character changes supporting the row oftarsal trichobothria (character 44), the branches of the tree in figure 26 are shown reduction of tarsal spines (character 3), and in table 4. the already mentioned paired spermathecae, Some traditional characters fit the tree well. are the synapomorphies of diplurines plus Among those are the characters traditionally Crassitarsae and Rastelloidina. Most ofthose used to define the non-atypoids. Platnick and characters are reversed or modified within Gerstch (1976) had proposed that the pres- Rastelloidina. 26 AMERICAN MUSEUM NOVITATES NO. 3056

TABLE 4 TABLE 4-(Continued) Synapomorphies (Asterisks [*] indicate character changes that oc- Taxon Changes cur in only some of the resolutions of clade 64) 41 (none, 0 > 1), Taxon Changes 67 (none, 3 > 0, 4 > 0) Mygalomorphae 24 (0 > 1), 46 (0 > 2), Ixamatus 22 (none*, 1 > 0), 49 (0 > 2), 51 (1 > 0), 24 (0 > 1) 38 (none, 0 > 2, 1 > 2), Xenonemesia 41(0> 1), 67 (3 > 0) 50(none,0> 1) Micromygale 6 (1 > 0), 7 (1 > 0), Atypidae 14(0> 1), 18 (0> 1), 9(1 > 0), 13 (2 > 0) 20(0> 1),47(0> 1), 50 (none, 0> 1), Microstigmata 30 (1 > 0), 50 (none, 1 > 2) 55 (O > 1) Pseudonemesia Antrod + Atypoides 3 (0> 1), 29 (0> 1) Ecuador 46(1 > 2) Aliatypus 23(0> 1) Mexico 32 (0 > 1), 38 (2 > 1) Mecicobothriidae 3(0> 1),4(1 >0), Nemesia 17*(none,0> 1), 14 (0 > 1), 29 (0 > 1) 40 (none*, 0 > 1), Scotinoecus 9 (none, 0 > 1), 57(0> 1) 36 (none, 0 > 1) Stenoterommata 40 (none, 0 > 1) Hexathele 44 (none, 4 > 0) Acanthogonatus 22 (1 > 0), Atrax 9 (none, 0 > 1), 40* (none, 1 > 0) 44 (none, 4 > 0), Heteromigas 30(none,0> 1) 67 (none, 1 > 2) Calathotarsus 41 (0 > 1), 67 (0 > 3), Porrhothele 3 (0 > 1), 9 (none, 1 > 0), 30 (none, 1 > 0) 34(0> 1), 44 (none, 0 > 4) Actinopus 59 (none, 0 > 1) Chilehexops Plesiolena 9 (1 > 0), 26 (1 > 0), 44 (1 > 4) Euagrus 67 (0 > 2) Idiops 30 (1 > 0), 51 (I > 0), Ischnothele 23 (0> 1), 69 (0> 1), 29* (none, 0 > 1), 67 (none, 0 > 5) 49 (none, 0 > 2) Neocteniza 6 (1 > 0), 27 (0 > 1), Diplura 29 (none, 0 > 1), 55 (0> 1), 63 (0> 1), 49 (none, 2 > 0), 50 (none, 1 > 0), 50* (none, 0 > 1) 67 (none, 5 > 0) Melloina Misgolas 3 (0 > 1), 11 (0 > 1), Paratropidinae 11(1 >0), 12(1 >0), 26 (1 > 0), 44 (1 > 0), 13 (4 > 3), 23 (0 > 1), 67 (none, 0> 5) 29(1 >0),42(1 >0), Ummidia 42 (0 > 1), 59 (none, 1 > 0) 43 (1 > 0), 49 (none, 1 > 2), Myrmekiaphila 50 (none, 0 > 2), Cyrtauchenius 67 (4 > 0) Bolostromus Ischnocolus 2(0> 1), 13(4> 1), 34(0> 1) Fufius 2 (0> 1), 57 (0> 1) Theraphosinae 29(1 >0) Rhytidicolus 3 (1 > 0), 44 (0 > 1), 19 (none, 0 > 1), Barychelidae 47 (0 > 1), 54 (1 > 0), 45 (none, 0> 1), 13 (none, 4 > 2) 49 (none, 1 > 2), Neodiplohele 13 (none, 2 > 4), 50* (none, 1 > 0), 40* (none, 0 > 1), 52 (none, 0 > 1) 1993 GOLOBOFF: REANALYSIS OF MYGALOMORPHS 27

TABLE 4-(Continued) TABLE 4-(Continued) Taxon Changes Taxon Changes

Clade 42 25 (1 > 2), 38 (2 > 1), 45 (none, 1 > 0), 61(1 > 2) 49 (none, 1 > 2), 52 (none, 1 > 0), Clade 43 16 (1 > 0), 17 (1 > 0), 53(1 39(1 >0),54(1 >0), >0),47(1 >0), 67 (3 > 0) 57 (0> 1), 68 (0> 1), 69(0> 1),70(0> 1) Clade 59 4 (1 > 2), 19 (none, 1 > 0), Clade 44 4(1 >2),45(0> 1), 34(0> 1), 63(0> 45 (none, 0> 1), 1),64(0> 1), 49 (none, 2> 30 (none, 1 > 0) 1), 52 (none, 0> 1) Clade 45 57(0> 1) Clade 60 12(0> 1), Clade 46 10(1 >0) 13 (none, 2 > 4), Clade 47 27(0 > 1), 52(0 > 1), 42 (0 > 1), 44 (0 > 2), 55 (0> 1), 67 (none, 3 > 0, 3 > 4) 30 (none, 1 > 0) Clade 61 11 (1 > 0), Clade 48 21 (1 > 0), 31 (0 > 1), 38 (none*, 1 > 2) 32 (0 > 1), 33 (0 > 1), Clade 62 2 (0 > 1), 40 (none, 0 > 1), 50 (none, 1 > 0), 65(0> 1) 56(0> 1), Clade 63 > > 67 (none, 0 > 5) 0 (1 0), 1(1 0), 16 (0> 1), Clade 49 3(1 >0),30(1 >0), 29* (none, 1 > 0), 38 (1 > 2), 50* (none, 0 > 1) 49 (none, 1 > 2), 62(0> 1), Clade 64 4(1 >0), 10(0> 1), 50 (none, 0 > 2), 22 (none*, 0> 1), 66(0> 1) 49 (none, 0> 1, 2> 1), 59 (none, 0 > 1) Clade 50 35(0> 1) Clade 65 Clade 51 2(0> 8 (1 > 2), 17 (none*, 0> 1), 1),24(0> 1) 19 (none, 0 > 1), Clade 52 3(1 >0), 11(1 >0), 47 (0> 1), 26(0> 1) 45 (none, 0> 1), Clade 53 4 (none, (0 > 1), 52 (none, 0 > 1) 13 (none, 2 > 4), Clade 66 2 (1 > 0), 17* (none, 0 > 1), 14 (0 > 1), 29* (none, 0 > 1), 15 (none, 0 > 1), 34(1 > 0), 36(1 > 0), 22(1 >0) 48(1 > 0), 60(1 > 0), Clade 54 39(1 > 2), 50 (none, 1 > 2) 49 (none, 0 > 1, 2 > 1), Clade 55 13 (2 > 3), 23 (0> 1), 59 (none, 0 > 1) 44(0> 1) Clade 67 7 (0 > 1), 23 (0 > 1), Clade 56 4 (none, 0 > 1), 36 (none, 1 > 0), 15 (none, 0 > 1), 67 (none, 1 > 2) 41 (none, 1 > 0), Clade 68 3 (0> 1), 9 (0> 1), 43 (0> 1), 11 (0 > 1), 13 (0 > 2), 67 (none, 0 > 4, 3 > 4) 44 (4 > 0), 46 (0> 1), Clade 57 37 (0 > 2), 50 (none*, 0> 1) 22* (none, 0 > 1) Clade 69 46 (0 > 1), 44 (none, 4 > 0) Clade 58 18(0> 1), Clade 70 6 (0> 1), 67 (0> 3), 19 (none, 0> 1), 29 (none, 0 > 1), 20(0> 1),22(0> 1), 49 (none, 0 > 2) 28 AMERICAN MUSEUM NOVITATES NO. 3056

TABLE 4-(Continued) pseudosegmented (character 29, 1.3), and the intercheliceral tumescence (character 41, 1.7); Taxon Changes most of them were already known to be of Clade 71 6 (0 > 1), 9 (none,0 > 1), limited use at higher levels. 15(0> 1), 67 (0> 1) The data set did not include any behavioral characters. An interesting implication of the Clade 72 7 (0> 1), 30 (0> 1), could be 34(0> 1), present results is that making webs 36(none,0> 1) a synapomorphy for all the mygalomorphs excluding Atypidae plus Antrodiaetidae (i.e., Clade 73 5 (0 > 1), 8 (none, 0 > 1), clade 76), secondarily lost in most mygalo- 21 (0 > 1), 25 (0 > 1), and Por- 28 (1 > 0), morphs. Scotinoecus (personal obs.) 36 (none, 0 > 1), rhothele, as well as mecicobothriids and di- 39 (0 > 1), 61 (0 > 1) plurids, build webs used in prey capture. Thus, the Rastelloidina (including the trapdoor spi- Clade 74 8 (0> 3), 9 (0> 1), In 17 (0> 1), ders), would be burrowers secondarily. 50 (none, 1 > 0), support for that idea, it can be mentioned 53 (none, 1 > 0) that the "cyrtaucheniid" Fufius (basal to the rastelloid clade) lives in silken tubes in crev- Clade 75 0 (0 > 1), 1 (none, 0 > 1), 2 (none, 0 > 1), ices, prolonged with silk (personal obs.), and 8 (none, 0 > 1), (as in some nemesiids, such as Acanthogon- 38 (none, 0 > 1, 2 > 1), atus) those constructs might represent re- 44 (none, 3 > 4), duced "webs." 48 (0> 1), 50 (none, 1 > 0), 58 (0 > 1), SOME SELF CRITICISM 53 (none, 0 > 1) Besides the ambiguity in the resolution of Clade 76 1 (none, 1 > 0), clade 64, there are several potential problems 2 (none, 1 > 0), with the above analysis. First, some alter- 16 (0 > 1), 19 (0 > 1), native topologies have a fit almost as good 60 (1 > 0), as that for the tree in figure 26, which is thus 38 (none, 1 > 0, 2 > 0), only slightly preferable to them. The area with 44 (none, 4 > 3) more character conflict seems to be the basal part of the tree, i.e., that part comprising the hexathelids and non-diplurine diplurids. For The cladograms obtained also reflect quite example, if Hexathelidae is made paraphy- well the developmental characters (charac- letic (by placing Euagrus as the sister group ters 59, 60); although the development of of Chilehexops, then moving clade 71 as the many taxa has not been studied, the opti- sister group of Porrhothele, and clade 68 as mization ofthose characters on the trees leads the sister group ofclade 67), the difference in to predictions that second instar larvae of total fit is only 0.3. That tree would sacrifice unstudied taxa in the Rastelloidina and Cras- two steps in character 15, and one in char- sitarsae (including Microstigmatidae) should acters 30 and 67, and would save one step in have a bent cephalothorax, and unstudied taxa characters 7, 9, 34, and 44. Both trees would in the Mecicobothriidae and Hexathelidae have the same length; but the former three should have the cephalothorax and abdomen characters show less homoplasy than the lat- in the same plane. Because there are many ter four, hence the total fit weakly favors the missing entries for these characters there is monophyly of Hexathelidae. some ambiguity in their optimization. Second, the analysis is based on only some Among the characters that fit the tree most taxa. Including more taxa may greatly change poorly are the tarsal spines (character 3, with the outcome (examples abound in the cla- a fit of 1.3), the reduction of the ITC teeth distic literature!); the only way to have a (character 9, 1.4), the domed apical article of cladogram guaranteed to be optimal when all the PLS (character 22, 1.7), the male tarsi the mygalomorph genera are considered is to 1993 GOLOBOFF: REANALYSIS OF MYGALOMORPHS 29 conduct a global parsimony analysis of all cladogram in figure 26 suggests indeed that those genera. Pee-Wee, the computer pro- most ofthe apparent "apomorphies" for some gram used here to search for trees, certainly microstigmatids are the plesiomorphic states could not handle a data set including the es- for the group, reversed in Micromygale. timated 250 mygalomorph genera, and, es- Among these reversals is the "plesiomorph- pecially, no arachnologist has yet been able ic" presence of ALS. The fittest trees I was to gather that huge amount ofinformation in able to find in which the presence of ALS in the form of a data matrix. This problem can Micromygale was a plesiomorphic retention be overcome by a judicious choice of taxa; if imply the polyphyly ofMicrostigmatidae and taxa including all the relevant character com- require a decrease of total fit of 10.9. Since binations are included in the analysis, the not allowing spinnerets to be regained would results are more likely to be stable under the imply disregarding other lines of evidence addition of new taxa. There are several taxa, solely to fit the preconception (which I myself not included in the present analysis, which share!) that gaining an extra pair ofspinnerets might introduce important modifications. is much rarer than losing it, the implications Among those are some Cyrtaucheniidae (a of the present data are followed and Micro- group which is basal to the Rastelloidina), mygale is considered as the most apomorphic and several "nemesiid" genera. (instead of the most plesiomorphic) repre- Third, for the results based on the weight- sentative of the Microstigmatidae. ing method used here to be correct, it is nec- NOMENCLATURAL IMPLICATIONS essary that the present data set adequately represents the relative amounts ofhomoplasy The results ofthe present study suggest that of the characters. I suspect that the weights the groups Fornicephalae and Tuberculotae assigned would not change much ifmore gen- will have to be abandoned. As suspected by era were included, but that could only be de- Platnick (1977), Atypoidea in the traditional termined by adding them. sense (i.e., that of Simon, 1892) is a para- Fourth, the characters that support many phyletic group. However, the group could be ofthe branches in the cladogram appear very made monophyletic just by excluding the weak. Table 4 lists the actual or potential Mecicobothriidae (and would then agree with synapomorphies for each clade in figure 26. the Atypoidea of Simon, 1903). Names for a Many of the characters can be optimized, few ofthe other higher taxa in the cladogram along some branches, in different ways (in have already been proposed by some arach- most cases, either as two gains or as a gain nologists. However, some of those names and subsequent loss), and they do not con- have been used by other workers to refer to stitute actual support for a hypothesis of different groups. For example, Aviculariidae monophyly. This problem is somewhat re- designated all mygalomorphs except atypids, lated to the previous two, in that the ambi- antrodiaetids, and mecicobothriids for Simon guity could be overcome by adding taxa. (1892), but included mecicobothriids for Fifth, there is a regain ofALS in Micromy- Simon (1903); Ctenizoidea corresponded to gale. Arachnologists have always been reluc- the Aviculariidae ofSimon (1892) for Cham- tant to pose multiple appearances of organs berlin and Ivie (1945), but to Ctenizidae, as complex as spinnerets or the cribellum (see Atypidae, Migidae, and Paratropididae for Platnick, 1977, for discussion) and have fa- Savory (1926). This makes it difficult to de- vored secondary losses. Raven (1985) has cide which choice would imply greater no- suggested that many ofthe apparent plesiom- menclatural stability. By using Avicularioi- orphies in Micromygale may be neotenically dea to refer to mecicobothriids plus the rest correlated with the small size and that the of non-atypoids (i.e., Aviculariidae sensu apparently "apomorphic" states in the other Simon, 1903), the oldest name can be pre- microstigmatids would actually be the orig- served. For the rest of the cladogram, Rav- inal (adult) characters, which might place mi- en's (1985) classification is the only one that crostigmatids close to nemesiids, or bary- contains roughly corresponding groups. Thus, chelids, or both. Whether or not neoteny is there are two options: either using Raven's the explanation for that phenomenon, the names with some relimitation, or creating 30 AMERICAN MUSEUM NOVITATES NO. 3056 completely new names. I prefer the first op- The family Nemesiidae probably will have tion, and propose to relimit both Orthopal- to be split in the future. Some genera previ- pae and Quadrithelina to include also the Mi- ously included in the Nemesiidae are more crostigmatidae and Rastelloidina (i.e., clades closely related to genera included in the Mi- 73 and 72, respectively), and to relimit Cras- crostigmatidae. Those are Xenonemesia, sitarsae to include the Microstigmatidae (i.e., Ixamatus, Xamiatus, and possibly the neo- clade 64). No author has so far proposed a tropical Spelocteniza Gertsch, which has group equivalent to clade 68; I propose the modified booklung openings, an elevated tar- name Bipectina for that group. sal organ (contra Raven, 1985: 111), basal At the family level, and given the problems and medial article ofthe PLS without spigots, discussed in the preceding section, the pres- and cuticle apparently scaly (the topotypical ent results are considered too preliminary to specimens studied here were not examined warrant nomenclatural changes. If future with SEM). Thus, either Nemesiidae must be studies support the hypotheses of relation- divided into several families or the Micro- ships proposed here, the family Cyrtauch- stigmatidae must become a subfamily ofNe- eniidae will require substantial changes in mesiidae. More taxa should be included in composition. Raven (1985: 65) had already the analysis to settle that question and allow suspected that the North American Eucten- an adequate rearrangement of the Nemesi- izinae could be more closely related to the idae. other Rastelloidina than to the other "cyr- taucheniids." Euctenizinae may well deserve REFERENCES familial status (studies on North America rastelloids by Platnick and Gertsch may clar- Carpenter, J. M. ify this point). Even if euctenizines are ex- 1988. Choosing among multiple equally par- cluded, however, Cyrtaucheniidae will still simonious cladograms. Cladistics 4: a doubtful group; although the "typical" 291-296. be Chamberlin, R. V., and W. 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