2001. The Journal of Arachnology 29:378±387

SYNONYMS OF FRONTINELLA TIBIALIS (ARANEAE, )

G. Ibarra-NuÂnÄez1, J. A. GarcõÂa1, M. L. JimeÂnez2 and A. MazarieÂgos1: 1El Colegio de la Frontera Sur. Carr. Antiguo Aeropuerto km 2.5, Apdo. Postal 36, Tapachula, Chiapas 30700, MeÂxico; 2Centro de Investigaciones BioloÂgicas del Noroeste S.C. Apartado Postal 128, La Paz, Baja California Sur, 2300, MeÂxico. email: gibarra@ tap-ecosur.edu.mx

ABSTRACT. Synonymy among three species of linyphid of the Frontinella F. O. Pickard- Cambridge 1902 is established based on ®eld association between males and females, mating records, and morphological data from recently collected specimens. It is concluded that F. caudata Gertsch & Davis 1946 and F. lepidula Gertsch & Davis 1946 are both junior synonyms of F. tibialis F. O. Pickard- Cambridge 1902. A redescription of this species is included.

RESUMEN. Se establece la sinonimia entre tres especies de aranÄas linõ®das del geÂnero Frontinella F. O. Pickard-Cambridge 1902, con base en datos de campo sobre la asociacioÂn entre machos y hembras, registros de apareamientos, y datos de la morfologõÂa de especimenes recientemente colectados. Se concluye que F. caudata Gertsch & Davis 1946 y F. lepidula Gertsch & Davis 1946 son sinoÂnimos junior de F. tibialis F. O. Pickard-Cambridge 1902. Se incluye una redescripcioÂn de esta especie. Keywords: Linyphiidae, Frontinella, synonymy, Mexico, Chiapas.

The genus Frontinella was created by ard-Cambridge 1902, F. tibialis F. O. Pickard- F. O. Pickard-Cambridge in 1902 for eight spe- Cambridge 1902, F. caudata Gertsch & Davis cies found in Central and North America. Pe- 1946, F. huachuca benevola Gertsch & Davis trunkevitch (1911) made Frontinella a junior 1946, F. lepidula Gertsch & Davis 1946 and F. synonym of Linyphia Latreille 1804 without ar- potosia Gertsch & Davis 1946. Only two of gument, but Blauvelt (1936) resurrected Fron- these have been recorded for the state of Chia- tinella from this synonymy when she made a pas, Mexico: F. caudata and F. lepidula revision of Linyphia and several related genera. (Gertsch & Davis 1946; Hoffmann 1976). Roewer (1942, 1954) and Bonnet (1956, 1957), In 1995, we collected female and male in their respective catalogues, included Fronti- specimens of Frontinella in coffee plantations nella as a junior synonym of Linyphia, but they in southeast Chiapas. The female specimens did not give any argument. Nevertheless, from were identi®ed as F. caudata, some males as that time on most authors considered Frontinel- F. tibialis, and other males as F. lepidula. la a valid genus name (Kaston 1938, 1948; Frontinella tibialis and F. lepidula were de- Gertsch & Jellison 1939; Muma 1943; Brignoli scribed from only one male specimen each, F. 1983; Millidge 1984; Platnick 1989, 1993, 1997; caudata was described from only female spec- Breene et al. 1993), and several new species imens. We found accompanying males in the were described (Gertsch & Davis 1946; Bryant webs of several F. caudata females. Most of 1948; Kraus 1955; Li & Song 1993). Millidge these were identi®ed as F. tibialis, one as F. (1991) transferred F. uncata F. O. Pickard-Cam- lepidula. The ®nding of these pairs suggests bridge 1902 to the genus Novafrontina Millidge synonymy. 1991. At present there are 15 species world- In their description of F. lepidula, Gertsch wide, two from China and 13 from the Ameri- & Davis (1946) considered it near to F. tibi- cas, with nine of these species reported from alis but pointed out some differences: ``This Mexico: F. communis (Hentz 1850), F. laeta (O. is a smaller species than tibialis Cambridge. Pickard-Cambridge 1898), F. bicuspis F. O . The embolus of the male palpus is shorter and Pickard-Cambridge 1902, F. rustica F. O. Pick- less strongly curved at the apex, and the pa-

378 IBARRA-NUÂ NÄ EZ ET AL.ÐSYNONYMS OF FRONTINELLA TIBIALIS 379 tella of the palpus is armed with a long curved Instituto de BiologõÂa, Universidad Nacional spine instead of a short spur.'' Some of our AutoÂnoma de MeÂxico), were also examined male specimens could not be assigned to any for comparative analyses. We examined the of these two species because they have a patellar macroseta of both palpi for each col- ``short spur'' in one patella and a ``long lected male, and made some SEM photo- spine'' in the other, showing another indica- graphs of them. Additional information about tion of possible synonymy. Thus, we decided the type specimens, not included in the orig- to collect more specimens of both sexes, to inal descriptions, was provided by Dr. N. I. study and clarify the of the three Platnick and by Mr. P. D. Hillyard. The de- described species. scriptions in the taxonomic section were based on the specimens collected in this work, with METHODS each measurement noted as average and range The collecting site was the coffee plantation of variation (minimum and maximum); all of the ``Campo AgrõÂcola Experimental Rosa- measurements are in millimeters. Abbrevia- rio Izapa'' of the Instituto Nacional de Inves- tions used in text: ALE: anterior lateral eyes; tigaciones Forestales, AgrõÂcolas y Pecuarias AME: anterior median eyes; PLE: posterior (INIFAP), Municipio of Tuxtla Chico, 20 km lateral eyes; PME: posterior median eyes. Fe- NNE of Tapachula, Chiapas, at 400 m eleva- I (II, III, IV): ®rst femora (second, third, tion. This place was selected due to the abun- fourth); Me-I (II, III, IV): ®rst metatarsi (sec- dance of Frontinella. The ®rst collecting pe- ond, third, fourth); Pa-I (II, III, IV): ®rst pa- riod was from September±November 1995, tellae (second, third, fourth); Ta-I (II, III, IV): and the second in January 1998. The spiders ®rst tarsi (second, third, fourth); Ti-I (II, III, were collected by visual search of the webs of IV): ®rst tibiae (second, third, fourth). Cy: adult and subadult females (as it is dif®cult to cymbium; PCy: paracymbium. differentiate these two age classes in the ®eld); additionally, some solitary males were RESULTS collected to have enough specimens for the We collected 76 individuals from 55 webs: taxonomic study. All individuals found on the 39 webs were occupied by only one individual same web were put in the same container and (solitaries), the remaining 16 webs contained preserved in 70% ethanol. In the laboratory 37 spiders, from two to ®ve individuals per each specimen was tentatively identi®ed and web (Table 1). Most accompanied adult fe- the age class and sex was determined. Speci- males shared the web with only one male, but mens were deposited at the ColeccioÂn de Ar- a few were found with several males. Most anÄas del Sureste de MeÂxico (ECOTA-AR, El sub-adult females were alone on their webs, Colegio de la Frontera Sur, Tapachula, Chia- only one was found with a male. Inexplicably, pas, Mexico), the American Museum of Nat- one sub-adult female and one juvenile were ural History, New York (AMNH), and The found each on the web of an adult female. Natural History Museum, London (NHM). Most males were found on the webs of adult To study the specimens, we considered the females, but two males were each on the web characters in the original description of each of a juvenile (Table 1). species. All adults (females and males) were In one web, one adult female was found measured to compare their variability in re- with four males (Table 2), and we observed lation to the measurements of the correspond- the end of copulation between one of these ing type as noted in the original descriptions. males and the female. On a date subsequent The measured characters were total length, to the sampling period, one of the authors (J. carapace length, carapace width, and length of A. GarcõÂa) observed another pair in a web, each segment of the ®rst leg from femur to copulating at least two times during the 30 tarsus. Specimens of both sexes were sent to min of observation. Dr. N. I. Platnick (AMNH), and to Mr. P. D. The measurements from 30 collected fe- Hillyard (NHM), for comparison with the cor- males showed that most mean values of these responding types in those museums. The par- females are slightly smaller than those of the atypes of F. caudata and of F. huachuca be- F. caudata holotype and paratype, but the var- nevola, deposited in the ColeccioÂn Nacional iation of these types are well inside the vari- de AraÂcnidos (CNAN, Dr.Tila MarõÂa PeÂrez, ation ranges of the collected females. Besides 380 THE JOURNAL OF ARACHNOLOGY

Table 1.ÐNumbers of specimens collected on each web, with their corresponding sex/age. F ϭ fe- male(s), M ϭ male(s).

Sub-adult No. in web Adult F F Adult M Juveniles totals Solitaries 18 14 7 39 with F 1 16 1 18 with 1 M 8 1 2 11 with Ͼ1M 3 3 with sub-adult F 1 1 2 with juveniles 1 2 3 Age/sex totals 31 16 26 3 76 the size variations, the study of the adult fe- We found males with ``long spines'' on males showed no major differences among both palpi, males with ``short spurs'' on both them, nor in respect to the characters noted in palpi, but also males with a ``short spur'' on the description of F. caudata, or those ob- one palp and a ``long spine'' on the other (Ta- served in the paratype of this species. The epi- ble 2). This mixed condition seems to indicate gyna were very similar for all females, para- that the macroseta is originally long, but it can type included, in respect to form and position break and lose its slender distal part, with only of the openings, dorsal and ventral plates (as the thick base remaining. SEM images de®ned by Millidge 1984). Although some showed that the ``short spur'' is a patellar ma- variability was recorded in the form and size croseta broken in the area where its diameter of the opisthosoma and its caudal tubercle, is reduced (Figs. 1, 2). this could be due to differences in nutritional Our data also show that both solitary and condition, level of development of ovaries, or courting males (in the web of a female) show natural variability. the three conditions. Likewise, one female Gertsch & Davis (1946) considered the specimen was accompanied with males show- length of the patellar macroseta, and length ing these three conditions (Table 2). Also, the and curvature of the embolus as distinctive male observed copulating with a female on a characteristics between F. tibialis and F. lep- date subsequent to the collect period had a idula. The collected male specimens showed broken macroseta in one palp and a complete only slight variability among them in embolus macroseta in the other. characteristics. On the contrary, for the length As for the females, the measurements of the of the patellar macroseta we found two states, collected male specimens showed also over- the ``short spur'' of Gertsch & Davis (as in lapping ranges of size among them, and with the ®gs. 7a and 7b of F. tibialis by F. O. Pick- the holotype of F. lepidula. It was also found ard-Cambridge 1902, table XL), or the ``long that the collected males have a mastidion on spine'' (as in ®g. 11 of F. lepidula by Gertsch each chelicera (one small laterally directed tu- & Davis 1946). bercle at the anterior proximal surface of the

Table 2.ÐNumbers of specimens collected alone or accompanied, and the pedipalp-patella macrosetae condition of the corresponding males. F ϭ female, M ϭ male(s), BM ϭ broken macroseta, CM ϭ complete macroseta. * The values with the same subscript letter corresponds to the same female specimen.

M with 1 CM M with 2 CM M with 2 BM and1BM Solitary M 2 2 3 1 F with 1 M 2 4 2

1 F with Ͼ 1M* 2b ϩ 1c 2a ϩ 1b ϩ 1c 1b 1 subadult F with 1 M 1 1 juvenile F with 1 M 2 Totals 8 12 6 IBARRA-NUÂ NÄ EZ ET AL.ÐSYNONYMS OF FRONTINELLA TIBIALIS 381 cheliceral base, just below the clypeus). The because of the variability of the opisthosomal mastidion size is variable, from sharply point- size in spiders (Blauvelt 1936; Hormiga ed in some specimens to blunt and very re- 1994a). It was not possible to obtain other duced in others. Some specimens have mas- measurements from this type. Nevertheless, tidia of different size, one pointed and one the maximum value of total length found from reduced. In some of the males, the chelicerae our specimens is only slightly below of that were so retracted that the mastidion was not noted for the type of F. tibialis (4.4 vs. 4.5). directly visible because it was covered by the Additionally, there is a high variability in size clypeus, but a careful examination showed its in the collected male specimens, where the presence in all collected males. smallest male is only one half of the largest (2.2 to 4.4). Furthermore, the range of varia- DISCUSSION tion for each measured character overlapped In some linyphiid species, the males re- among the three variants of the collected spond to a species-speci®c sex pheromone males (with 2 complete macroseta, with 2 bro- present in the silk of adult female webs by ken macroseta, and with one complete and approaching the female and initiating court- one broken macroseta), and the measurements ship behavior; in some cases the males can of the F. lepidula holotype were also inside stay in the web for some time, and even cop- these variation ranges. As F. tibialis was de- ulate several times with the female (Rovner scribed from only one specimen, there were 1968; Austad 1982; Suter & Renkes 1982; no records about its size variability. Thus, the Watson 1986, 1995; Wiley-Robertson & Adler slight difference in size does not contradict the 1994). The presence of the males in the fe- conspeci®ty of our specimens with the type of males' webs, and the copulations observed F. tibialis. constitute sound evidence of conspeci®city, Concerning the differences between F. tib- especially since no other related species were ialis and F. lepidula in size and curvature of found at the collecting site. the embolus mentioned by Gertsch & Davis The morphological similarity between the (1946), we consider these as minor differences collected females and the paratype of F. cau- in comparison with other Frontinella species data (especially with regard to the epigynum), (judging from the drawings of F. O. Pickard- and the fact that the measurements of the ho- Cambridge 1902; Blauvelt 1936; Gertsch & lotype and paratype of this species are inside Davis 1946; and Song et al. 1999), where the the ranges of the collected females, indicate pedipalpal bulbs (subtegulum, tegulum, em- that F. caudata and the collected specimens bolic division, particularly the embolus and la- are the same species. mella characteristica) are conspicuously dif- Platnick (pers. comm.) considered a female ferent among species. When the palpal bulb specimen sent to him as the same species as of the collected specimens is observed from the F. caudata holotype, and the male speci- different angles the embolus becomes more or mens sent to him as the same species of the less curved, and the lamella characteristica be- F. lepidula holotype. For the male specimens comes more or less wide. Additionally, the sent to be compared with F. tibialis, Hillyard type of F. tibialis comes from a locality near (pers. comm.) considered that ``the pedipalp the Gulf of Mexico coast, but the type of F. macroseta is not signi®cantly different'', but lepidula and the males we collected come that ``there is some doubt that these two spe- from a locality near the Paci®c coast of Mex- cies are conspeci®c'', because he noted some ico; therefore, it is possible that these are op- differences such as total length (``the type is posite ends of a geographic variability spec- slightly larger''), the size of the palp (``more trum concerning the size and curvature of the robust in the type''), the shape of the embolus embolus. (broader and more curved at its tip), and the F. O. Pickard-Cambridge (1902) noted ex- presence of mastidia (``the type does not have plicitly in his key to Frontinella species that a single mastidion on the basal segment of F. tibialis lacks mastidia, but all our male each chelicera''). specimens have one on each chelicera, al- In the original description of F. tibialis, F. though sometimes a very reduced one. The O. Pickard-Cambridge (1902) noted only the presence of mastidia was not mentioned by total length, which is not a reliable character Gertsch & Davis (1946), but Platnick (pers. 382 THE JOURNAL OF ARACHNOLOGY comm.) con®rmed its presence in the F. lep- TAXONOMY idula holotype. Most species of Frontinella Figs. 1±10 with known males have a mastidion on their Frontinella tibialis chelicerae: F. laeta and F. bicuspis (F. O. F. O. Pickard-Cambridge 1902 Pickard-Cambridge 1902), F. communis Frontinella tibialis F. O. Pickard-Cambridge, 1902: (Blauvelt 1936), F. huachuca and F. huachu- 422, plate XL ®gs. 7a±b (; Gertsch & Davis, ca benevola (Platnick pers. comm. and per- 1946: 3. sonal observation of the male paratype of F. Linyphia tibialis, Petrunkevitch 1911: 255; Roewer huachuca benevola in the CNAN). Thus, the 1942: 591; Bonnet 1957: 2531. presence of mastidia seems not to be rare in Frontinella caudata Gertsch & Davis, 1946: 4, ®g. 6 &; Brignoli 1983: 294. NEW SYNONYMY. this genus. The collected male specimens Frontinella lepidula Gertsch & Davis, 1946: 4±5, showed a high variability in the size of their ®gs. 10±11 (; Brignoli 1983: 294. NEW SYN- mastidia. As F. tibialis was described from ONYMY. only one specimen, there were no records about mastidion variability. Also, it is possible Types.ÐMale holotype of F. tibialis from that this difference corresponds to a spectrum Teapa, Tabasco, Mexico, in the collection of of variability between the two populations Goodman & Salvin, deposited in NHM (not (Gulf coast and Paci®c coast). examined). Female holotype of F. caudata Gertsch & Davis (1946) said about Fron- from Chilpancingo, Guerrero, Mexico, depos- tinella, ``It is notable that the males of the ited in the AMNH, female paratypes from three new species herein described all have Chilpancingo, Guerrero, Mapastepec and Ta- the patella of the palpus set with a long dorsal pachula, Chiapas, Mexico, deposited in the spine. In all the other known species this spine AMNH and in the CNAN (examined). Male is modi®ed into a short spur.'' As both spe- holotype of F. lepidula from Tonala, Chiapas, Mexico, deposited in the AMNH (not exam- cies, F. tibialis and F. lepidula, were de- ined). scribed each with only one specimen, the var- Diagnosis.ÐThis is a tentative diagnosis, iation existing in this character was not because several species of this genus are observed. We do not know when and how the known only from one sex, and we did not re- patellar macroseta breaks. As they are present vise this genus. A possible autapomorphy of only in the males, it is possible that these this species is the distinctive form of the la- structures are related to reproductive activi- mella characteristica in the male pedipalp ties. It would be necessary to study the repro- (Figs. 4, 7), clearly different from that of all ductive behavior of this species to know more other known males. The tibia's relative length, about the function of this structure. twice as long as the patella in the male pedi- Other similarities that support the hypoth- palp (Fig. 3), separates this species from most esis of conspeci®ty between the type of F. tib- others species with known males (with tibiae ialis and the collected male specimens are the less than twice the patella length), except from presence of macrosetae in the mesal border of F. potosia, but in F. potosia the patellar ma- the cymbium (Fig. 7), the relative size be- croseta of the pedipalp does not have a wid- tween pedipalp's patella and the tibia (tibia ened base as in F. tibialis. The position of the about twice as long as patella), the form of copulatory openings (on the lateral borders, the pedipalp's tibia (widening to its distal end, about at the middle of the distance between Figs. 6±8), and the form of the male sternum, the anterior and posterior borders of the dorsal narrowly produced between coxae IV (as in plate, Fig. 9) distinguish this species from table XL, ®g. 7 of F. O. Pickard-Cambridge most other species with known females (hav- 1902). ing the copulatory openings on the anterior From this evidence, we conclude that F. border of the dorsal plate), except from F. zhui caudata and F. lepidula are junior synonyms Li & Song 1993, but in this species the pos- of F. tibialis by the principle of priority (Ar- terior border of the dorsal plate is notoriously ticle 23 of the ICZN). As the original descrip- rounded and extended backwards. tion of F. tibialis is very short, we include Description.ÐMale: (n ϭ 26) Total length here a redescription of this species based on 3.25 (2.16±4.43), carapace length 1.49 (0.97± the specimens collected in this work. 1.86), carapace width 0.98 (0.70±1.20). Di- IBARRA-NUÂ NÄ EZ ET AL.ÐSYNONYMS OF FRONTINELLA TIBIALIS 383

Figures 1±6.ÐFrontinella tibialis, scanning electron micrographs of male pedipalp structures: 1. Com- plete macroseta of patella; 2. Broken macroseta of patella showing fracture point; 3. Row of setigerous cusps on mesal face of right femur, with enlargement of one cusp (inset) (scale bar for femur 50.0 ␮m); 4. Ectoventral view of left pedipalp showing visible parts in unexpanded condition; 5. Ectodorsal view of right pedipalp in unexpanded condition showing paracymbium with setae (white arrow) and membrane of cymbium (black arrow) (scale bar 46.5 ␮m); 6. Dorsal view of right tibia showing three retrolateral (white arrows) and two prolateral (black arrows) trichobothria (scale bar ϭ 29.4 ␮m). Abbreviations: Cy ϭ cymbium; E ϭ embolus; EM ϭ embolic membrane; Fe ϭ femur; LC ϭ lamella characteristica; Pa ϭ patella; Pcy ϭ paracymbium; Sp ϭ spine of lamella characteristica; SPT ϭ suprategulum; ST ϭ subte- gulum; T ϭ tegulum; Ti ϭ tibia. 384 THE JOURNAL OF ARACHNOLOGY

Figures 7±10.ÐFrontinella tibialis, drawings of genitalia. 7±8. Left male pedipalp with expanded bulb. 7. Meso-ventral view; 8. Ecto-dorsal view; 9±10. Female epigynum. 9. Ventral view; 10. Dorsal view of cleared epigynum. Scale bars ϭ 0.1 mm. Abbreviations: Pedipalp: BH ϭ basal hematodocha; Cl ϭ column; Cy ϭ cymbium; CyM ϭ cymbium macrosetae; E ϭ embolus; ED ϭ ejaculatory duct; EM ϭ embolic membrane; Fu: ϭ fundus; LC ϭ lamella characteristica; LP ϭ lateral process of lamella characteristica; M ϭ membrane; Pcy ϭ paracymbium; Pe ϭ petiole; SPT ϭ suprategulum; ST ϭ subtegulum; T ϭ tegulum; Ti ϭ tibia. Epigynum: A ϭ atrium; CD ϭ copulatory duct; CO ϭ copulatory opening; DP ϭ dorsal plate; FD ϭ fertilization duct; S ϭ spermatheca; VP ϭ ventral plate. ameter of AME 0.08 (0.07±0.09), ALE 0.09 (0.30±0.50), Ti-I 2.03 (1.40±2.63), Me-I 2.33 (0.07±0.11), PME 0.09 (0.08±0.09), PLE 0.08 (1.60±3.26), Ta-I 1.23 (0.90±1.53), Pa-II 0.35 (0.07±0.11). Separation between AME 0.06 (0.28±0.43), Ti-II 1.56 (1.08±2.03), Pa-III (0.05±0.07), PME 0.08 (0.07±0.11), AME and 0.28 (0.20±0.37), Ti-III 0.81 (0.54±1.07), Pa- ALE 0.11 (0.08±0.13), PME and PLE 0.13 IV 0.35 (0.24±0.47), Ti-IV 1.42 (0.94±2.00). (0.09±0.15). Clypeus height 0.25 (0.19±0.32). Carapace with a distinct but shallow tho- Length of Fe-I 2.38 (1.60±3.16), Pa-I 0.39 racic groove. Eyes on low tubercles, anterior IBARRA-NUÂ NÄ EZ ET AL.ÐSYNONYMS OF FRONTINELLA TIBIALIS 385 eye row moderately recurved, posterior eye on its apex, more or less parallel to Cy, em- row slightly recurved, lateral eyes contiguous. bolic membrane and membrane both parallel Labium about 1.5 times wider than long, in to embolus, ending as membranous strips that close contact with sternum. Sternum scuti- touch the embolus tip (Figs. 4, 5, 7 & 8). form, produced behind between coxae IV. Carapace almost glabrous, sternum and en- Cheliceral base with a narrow but distinct dites with sparse setae. Chelicerae with sparse stridulatory band on the ectal side; with a short setae on anterior face, apex of outer mastidion on the anterior face near to its base, sides and a few setae bordering both cheliceral varying from a sharply pointed tubercle to a margins. Legs with scattered setae, more nu- reduced blunt one; anterior face of chelicerae merous and stiff on Me and Ta; stiff setae on with scattered setigerous cusps. Chelicerae setigerous cusps of Fe. Bristles on legs as fol- with 4±5 promarginal teeth and 4±5 small re- lows: 2 dorsal on Pa and Ti-I to IV; 1 ventral tromarginal teeth. Endites with a diagonal ca- on Ti-I and II; 1 prolateral on Ti-I; 1 retrola- rina on the outer half of the distal border. Legs teral on Ti-I and II; 1 dorsal and 1 ventral on I-IIഠIV-III. Coxae IV separated by about one Me-III; 1 dorsal on Me-IV. Opisthosoma with half of their width; Fe-I to IV with a few lon- sparse, short setae and with two groups of gitudinal ventral and lateral series of setiger- long bristles on its anterior end, above each ous cusps, more conspicuous on Fe-I and II. side of the pedicel. Opisthosoma elongated-oval from above, Pedipalp: with sparse setae form femur to about twice as long as wide; more or less rect- Cy; patella with a long distal dorsal macroseta angular in lateral view, with a scarcely devel- on dorso-distal protuberance, the macroseta oped rounded caudal tubercle. proximal ¼ is thick and curved to the outer Pedipalp: Femur with small setigerous side (forming an angle of about 90º from the cusps on its dorsal, and ectal faces; mesal face femur axis), and the rest thin and more or less smooth with only 5±6 setigerous cusps in a straight, tapering to a point, (Figs. 1, 2). Tibia longitudinal line (Fig. 3). Patella with a dorso- with longer setae forming an incomplete ring distal protuberance that supports a macroseta. near its distal border, with 2±3 retrolateral tri- Tibia about twice as long as patella (Fig. 3), chobothria and 1±2 prolateral trichobothria and widening to its distal end (Figs. 6±8). Cy (Fig. 6). Cy with 3±4 short thick macrosetae elongated, about twice as long as maximum on the distal half of its mesal border (Fig. 7). wide, narrowing toward its tip, with a trans- PCy with a few small setae on its distal half, lucent, convex membrane on the proximal half visible only at high magni®cation (Fig. 5). of its ectal border; alveolus occupying almost Coloration: Variations observed on both all proximal face, leaving unoccupied about fresh and older preserved specimens. Cara- distal one tenth (Figs. 4±5, 7±8). PCy a small pace, chelicerae and endites orange-brown, curved strip (intersegmental sensu Hormiga pars thoracica with faint dusky radiating lines, 1994b), similar in texture and coloration to eye tubercles black. Endites becoming white- Cy, about one sixth the Cy length, and touch- yellow towards their tip, with a black carina ing the Cy membrane (Figs. 5, 8). Subtegulum on the outer half of the distal border. Sternum transverse; tegulum trapezoidal, narrow on its and labium dusky orange-brown, with borders distal border; suprategulum visible distal to te- in front of endites and rear point infuscated. gulum, between the embolic division and the Pedipalpi dusky light-green to dark orange- alveolus (Figs. 4, 5). Embolic division con- brown. Legs with coxae to basal two thirds of nected to tegulum by a membranous column. femora light orange-brown, the rest dusky Lamella characteristica elongated, proximally light-green, darker from tibiae to tarsi. Opis- directed in the not expanded bulb, pointed on thosoma creamy-gray with dorsal orange- its proximal tip and reaching the base of Cy brown tinge, and sides with a dark band and (Figs. 4); with one short lateral process on its patches. Venter, caudal tubercle and spinnerets mesal side (Figs. 7, 8), that reaches the middle darker. of the mesal border of Cy, and with an incon- Female: (n ϭ 30) Total length 5.69 (4.46± spicuous spine (visible only at high magni®- 6.95), carapace length 1.96 (1.47±2.25), car- cation, Fig. 4) on the opposite side (the spur apace width 1.29 (0.87±1.67). Diameter of of the lamella in Blauvelt's 1936 description AME 0.09 (0.08±0.11), ALE 0.12 (0.11± of F. communis). Embolus pointed and curved 0.13), PME 0.11 (0.09±0.12), PLE 0.10 386 THE JOURNAL OF ARACHNOLOGY

(0.09±0.11). Separation between AME 0.06 to the posterior border of the epigynum, in (0.05±0.08), PME 0.09 (0.08±0.11), AME and contact with the border between dorsal plate ALE 0.16 (0.13±0.17), PME and PLE 0.16 and ventral plate, and curving dorsally at the (0.11±0.17). Clypeus height 0.25 (0.20±0.29). dorsal border of the genital opening (Fig. 10). Length of Fe-I 3.14 (2.56±3.56), Pa-I 0.63 Distribution.ÐMEXICO: Veracruz (Po- (0.50±0.70), Ti-I 2.91 (2.60±3.30), Me-I 3.11 trero), Tabasco (Teapa), Guerrero (Chilpan- (2.40±3.55), Ta-I 1.53 (1.13±1.80), Pa-II 0.60 cingo), and Chiapas (Tonala, Mapastepec, Ta- (0.53±0.63), Ti-II 2.25 (1.97±2.43), Pa-III pachula and Tuxtla Chico). 0.49 (0.43±0.50), Ti-III 1.26 (1.07±1.37), Pa- IV 0.55 (0.50±0.57), Ti-IV 2.20 (1.90±2.40). ACKNOWLEDGMENTS Female similar to male except in the follow- We thank the following persons: the au- ing characters: labium not in close contact with thorities of the Campo AgrõÂcola Experimental sternum. Cheliceral base clearly thickened Rosario Izapa, (INIFAP) for permitting us to proximally, without mastidia nor setigerous collect in their coffee plantation; T. M. PeÂrez cusps. Tarsi of pedipalpi with one simple claw. (CNAN, Instituto de BiologõÂa UNAM) for Legs I-IV-II-III. Fe without series of setigerous permitting us to examine the paratypes of F. cusps. Opisthosoma more or less trapezoidal in caudata and F. huachuca benevola; G. Nieto lateral view, with the rear side higher than the (El Colegio de la Frontera Sur) for her assis- anterior side, and with a pronounced caudal tu- tance with the scanning micrographs; N.I. bercle projected beyond the spinnerets. Opis- Platnick (AMNH) and P.D. Hillyard (NHM) thosoma with sparse short setae on the ventral kindly agreed to compare the specimens col- plate of epigynum. Carapace and chelicerae lected with the types deposited in their re- brown to dark brown. Sternum, labium and en- spective institutions; N. I. Platnick and G. dites dusky brown. Pedipalpi dusky light- Hormiga (George Washington University) and green, darker to the tarsi. Legs with distal half L. Leibensperger (Museum of Comparative of Fe light orange-brown. All Fe with a trans- Zoology) kindly provided useful information verse dark gray band on the distal ventral bor- and important literature. G. Hormiga, M. L. der. Opisthosoma dark brown to black, some Draney, J. Miller and the editors of the Journal specimens with a pair of small creamy white of Arachnology made many useful sugges- points on the middle of dorsum. Dorsum mar- tions for the improvement of the manuscript. gined with an irregular creamy white band in- This work was supported in part by a grant cluding the caudal tubercle, incomplete in from the Consejo Nacional de Ciencia y Tec- some specimens. Sides with another irregular nologõÂa, MeÂxico (CONACYT R28867-N). creamy white band at mid-height, and with four transversal (dorso-ventral) irregular dis- LITERATURE CITED continuous creamy white bands on the poste- Austad, S.N. 1982. First male sperm priority in the rior half. With a diffuse patch of creamy white , Frontinella pyramitela just above the anal tubercle. (Walckenaer). Evolution 36 (4):777±785. Epigynum (Figs. 9, 10) wider than long. Blauvelt, H.H. 1936. The comparative morphology of the secondary sexual organs of Linyphia and Ventral plate slightly convex, protruding very some related genera, including a revision of the little from the abdominal wall. Dorsal plate group. Vol. 2, pp. 81±171, In Festschrift zum 60 about as wide as long, concave in its anterior Geburtstage von Professor Dr. Embrik Strand. half forming an epigynal atrium where are Riga. found the exposed rounded copulatory open- Bonnet, P. 1956. Bibliographia Araneorum. Tome ings at each side, touching the border with the II, 2eme partie: C-F. Toulouse, Les Artisans de ventral plate (Fig. 9). In dorsal view (Fig. 10) L'Imprimerie Douladoure. Pp. 919±1926. copulatory ducts straight and short, pointing Bonnet, P. 1957. Bibliographia Araneorum. Tome to the sides, and leading directly to the sper- II, 3eme partie: G-M. Toulouse, Les Artisans de mathecae which are curved, kidney shaped. L'Imprimerie Douladoure. Pp. 1927±3026. Breene, R.G., D.A. Dean, M. Nyffeler & G.B. Ed- Fertilization ducts thin, long, leaving sper- wards. 1993. Biology, predation ecology and sig- mathecae from the internal curvature to the ni®cance of spiders in Texas cotton ecosystems midline, then making a loop around copula- with a key to the species. Texas Agriculture Ex- tory ducts, very near to the copulatory open- periment Station, College Station (B-1711). 115 ings, and then continuing more or less straight pp. IBARRA-NUÂ NÄ EZ ET AL.ÐSYNONYMS OF FRONTINELLA TIBIALIS 387

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