Introduction

The family Phrurolithidae comprises 25 genera and 404 species globally (World Spider Catalog, 2024). Approximately half of all Phrurolithidae species have been described within the past five years, primarily from Asia and North America, this last is a region recognized as a major center of diversity despite being considerably neglected (Platnick, 2019; World Spider Catalog, 2024). As with many other spider groups, most information derives from original taxonomic descriptions, which generally lack natural history data. Consequently, knowledge of this family's natural history is sparse and scattered throughout the literature. These spiders are known to be very small, ground-dwelling species inhabiting microhabitats such as under stones, fallen wood, and leaf litter (Dondale & Redner, 1982; Kaston, 1948). Females construct a scale-like egg sac, which is attached to stones, some abandon their egg sacs after completion while others guard them until hatching (Dondale & Redner, 1982; Platnick, 2019). Laboratory observations indicate that phrurolithids have a prey preference for springtails (Angulo-Ordoñes et al., 2019; Pekár & Jarab, 2011). Until recently, the mating behavior of Phrurolithidae remained virtually unknown, with Mu et al. (2022) providing the first brief description. The limited information available on phrurolithids is likely attributable to their small size and cryptic coloration.

Phonotimpus Gertsch & Davis, 1940, a Mexican genus of Phrurolithidae, comprises 32 species, each characterized by a restricted geographic distribution (Platnick et al., 2022; World Spider Catalog, 2024). Phonotimpus pennimaniChamé-Vázquez, Ibarra-Núñez & Jiménez, 2018 and P. talquian Chamé-Vázquez, Ibarra-Núñez & Jiménez, 2018 are two small-sized spiders (1.9-2.7 and 2.3-3.3 mm total length, respectively), highly abundant components of soil spider assemblages (Chamé-Vázquez et al., 2018). Phonotimpus pennimani inhabits leaf litter of shade coffee plantations at 920 m a.s.l., while P. talquian inhabits leaf litter of cloud forest at 2010 m a.s.l. on the slopes of Tacaná Volcano, Chiapas, Mexico (Chamé-Vázquez et al., 2018). Despite the abundance of both species (Chamé-Vázquez, 2011, 2015), no basic natural history information is available for either, or for any Phonotimpus species. This study presents laboratory and field observations on the natural history of both Phonotimpus species, encompassing prey capture, egg sac morphology, fecundity, maternal care behaviors, and molting retreats. Additionally, we report an interaction between a parasitoid wasp and P. pennimani. Furthermore, this study was part of a broader effort aimed at contributing to the taxonomy and phenology of both species.

Materials and methods

Most observations were conducted under laboratory conditions, with additional field observations made during sampling events. All spiders were collected from the type localities of P. pennimani (Alpujarras, 15.066878°N, 92.165833°W) and P. talquian (Talquián, 15.0875°N, 92.0989°W), both situated on the slopes of Tacaná Volcano in Chiapas, Mexico. Spiders were collected monthly by hand from March 2016 to October 2019 by dispersing leaf litter onto a white fabric sheet and capturing individuals using an aspirator. Voucher specimens are deposited at the Colección de Arácnidos del Sureste de México (ECOTAAR), El Colegio de la Frontera Sur, Tapachula, Chiapas, Mexico.

In the laboratory, each spider was housed individually in a 9 x 9 x 7 cm plastic container filled with a moistened plaster of Paris and activated charcoal substrate. While initially provided with dried leaves for shelter, these were replaced with translucent plastic straws due to a lack of observable behavior. Spiders were initially fed with Collembola, Psocoptera, and Formicidae. However, after observing a preference for springtails (Collembola), a diet of cultured springtails was provided twice per week. This diet consisted of Proisotoma sp. (Isotomidae), Pseudosinella sp., Lepidocyrtus sp. (Entomobryidae), and Cyphoderus sp. (Cyphoderidae). All spiders were maintained under laboratory conditions at 26 ± 1.3°C, 57 ± 4.3% relative humidity, with a 12:12 light:dark photoperiod, which corresponds to most of the year in this region. Further details on laboratory rearing can be found in Angulo-Ordoñez et al. (2019).

We adopted the life cycle terminology of Alvarado-Castro and Jiménez (2016) with the following modifications. A "juvenile" was defined as a spider from the first instar to the antepenultimate stage. The "penultimate" instar referred to the stage immediately preceding the adult molt. Penultimate males were identified by swollen tarsi lacking sclerotized structures, while penultimate females exhibited a slightly sclerotized gonopore region but lacked a vulva. An "adult male" possessed modified pedipalps (sclerotized copulatory bulb with discernible sclerites, and palpal tibia with apophyses), and an "adult female" had a distinct epigynum and vulva. Developmental stage determination was based on external morphology.

The shape, number of eggs, number of egg sacs per female, and embryonic period of egg sacs were estimated for females captured in the field with egg sacs and for females that produced egg sacs in the laboratory. We recorded the presence or absence of retreats and the molting dates of juveniles and penultimate individuals reared in laboratory. A total of 34 males, 56 females (including four with egg sacs collected from the field), eight penultimate males, five penultimate females, and 50 juveniles (plus 30 juveniles hatched in the laboratory) of P. pennimani were reared (Fig. 1 A-C). For P. talquian (Fig. 2 A-C), 11 males, 22 females, 15 penultimate males, seven penultimate females, and 60 juveniles were reared.

Figure 1
A-L. Phonotimpus pennimani. A. Female habitus. B. Male habitus. C. Penultimate male habitus. D-F. Female guarding egg sac. G. Exuvia and retreat. H-I. Molted juvenile with retreat and exuvia. J. Female consuming a springtail. K. Male consuming a springtail. L. Juvenile (first instar) consuming a springtail.

Figure 2
A-J. Phonotimpus talquian. A. Female habitus. B. Male habitus. C. Juvenile habitus. D. Female guarding egg sac. E. Female consuming a springtail. F. Male consuming a springtail. G. Juvenile molting. H. Exuvia and retreat. I-J. Molted juveniles with retreat and exuvia.

Results

Prey capture. In the field, we observed and captured a female of P. pennimani (21 February 2019) consuming an unidentified springtail of the family Tomoceridae (Collembola). Additionally, we collected a female of P. talquian (26 March 2019) ingesting a Pogonognathellus sp. (Tomoceridae) springtail. In the laboratory, juvenile, penultimate, and adult individuals of both species preyed on springtails that wandered nearby (Figs. 1J-K, 2E-F). Capture involved a short jump (less than one leg span) followed by subduing the prey with the forelegs and subsequently with the chelicerae. Even while guarding their egg sacs, P. pennimani females consumed springtails. In one instance, a juvenile of P. pennimani with the ventral macrosetae of its forelegs raised for a moment attacked a springtail larger than itself (Fig. 1 L). The springtail attempted to escape, jumped two times with the spider grasping it, but the escape mechanism was not effective, and the spider subdued its prey.

Retreat for molting in laboratory conditions. Juvenile and penultimate individuals of both species constructed a silk retreat for the molting process. These retreats were found at the top of the container, within the straws, or on the bare substratum. The retreats were variable (compare Figs. 1G-I, 2H-J) but typically consisted of two layers. The basal layer was generally more densely woven than the upper layer, with silk threads evident in both layers. During growth, some individuals constructed a retreat for one molting event but not for the subsequent ones. When juveniles did not build a retreat, they adhered to the top or walls of the container using their legs and molted (Fig. 2 G). No silk was found upon subsequent inspection. Several juveniles died during or immediately after the molting process. In one instance, a juvenile was unable to release its legs and became trapped in the exuvia, resulting in death.

Eggs sac, clutch size, and embryonic period. In the field, we observed four P. pennimani females with egg sac laid in dried leaves. The egg sacs were disk-like, with a hue ranging from pinkish to whitish. Under laboratory conditions, 21 P. pennimani females constructed their eggs sacs on the bare substratum, within straw or in container corners, while one P. talquian female laid her egg sac on a dried leaf (Figs. 1D-F, 2D, 3A). All egg sacs constructed under laboratory conditions were also disk-like, resembling those observed in the field, and composed of two layers: the upper layer appeared more densely woven than the basal one, with eggs packed in the center (Fig. 3 B-E). Neither species spun an extra layer of silk to protect themselves or the egg sacs. Twenty-one P. pennimani females built at least one egg sac, nine constructed a second, three built a third one, and one produced a fourth, whereas the single P. talquian female did not constructed other egg sac after the first one.

Figure 3
A, D-E. Egg sac of P. pennimani. B-C. Scanning electron micrograph (SEM) of egg sac upper layer detail. F. Eggs at different stages of embryonic development. G-H. Egg sac with parasitoid wasp Idris sp. I. Parasitoid wasp with puparium. J-L. Details of female parasitoid wasp of Idris sp.

For P. pennimani, the mean egg sac diameter was 5.62 mm ± 0.69 SD (n = 16), the mean number of eggs per sac was 8.93 eggs ± 2.64 SD (n = 16), with a range of six to 15 eggs, and the mean egg diameter was 0.49 mm ± 0.09 SD (n = 14). The mean embryonic development time, from oviposition to spiderling emergence, was 22.66 days ± 4.72 SD (n = 3 egg sacs). Furthermore, the female was not essential for offspring emergence; in two instances, offspring emerged from the egg sac after the females died. In a related case, a female built and guarded a second egg sac two days before the first clutch hatched and subsequently constructed a third egg sac two days before the second clutch emerged. We were unable to measure or count the eggs in the single P. talquian egg sac due to its loss before the offspring emergence.

Maternal care. In the field, we observed four P. pennimani females positioned themselves above their egg sacs. Under laboratory conditions, 21 P. pennimani females and one P. talquian female that constructed egg sacs exhibited maternal care by guarding their egg sacs. Females positioned themselves above their egg sacs, as observed in the field, or maintained contact using at least one leg (Figs. 1D-F, 2D). When disturbed during feeding, females rapidly sought refuge but subsequently returned to their egg sacs after several minutes, hours, or in some cases, not at all.

Endoparasitoids. Two female P. pennimani guarding their egg sacs were collected in the field (18 April 2018 and 14 January 2019). Each egg sac was attached to a dried leaf, and neither exhibited visible perforations or damage to the upper surface upon inspection. From both egg sacs emerged Idris sp. wasps (Scelionidae: Scelioninae; Fig. 3 J-L). A male wasp emerged from the first egg sac, leaving a circular opening in the upper layer. Inside the egg sac, we found five dead juveniles, one dead female wasp, and two wasp puparia (Fig. 3 G-H). The second egg sac contained an intact puparium, a male wasp (Fig. 3I), seven dead juveniles, and two unfertilized spider eggs.

Discussion

Prey capture. Here, we provide the first anecdotal field observation of P. pennimani and P. talquian preying on Tomoceridae springtails. Recently, Angulo-Ordoñez et al. (2019) described the predatory behavior of P. pennimani and P. talquian against four prey types under laboratory conditions. Their results indicated a preference for Collembola (springtails) over Hemiptera or Psocoptera, with both species employing ambush or active searching strategies. Pekár and Jarab (2011) observed broad prey acceptance in laboratory conditions for two phrurolithids, Liophrurillus flavitarsis (Lucas, 1846) and Phrurolithus festivus (C. L. Koch, 1835). Both species were more successful at capturing Sinella curviseta Brook, 1882, an Entomobryidae springtail, than fruit flies or aphids, and preferred prey smaller than their body size (Pekár & Jarab, 2011).

In our observations, both juveniles P. pennimani and P. talquian were capable of subduing large springtails, and no evidence of silk use for prey capture was observed, neither in the field nor under laboratory conditions. The long, ventral, and movable macrosetae on the tibia and metatarsi of the forelegs of both species appear to be adaptations for prey capture. Triaeris stenaspis Simon, 1892, a goblin spider (Oonopidae), resembles both P. pennimani and P. talquian in size and in possessing leg macrosetae on the tibia. Similarly, it does not construct a web for prey capture but instead employs a grasp-and-hold tactic (Korenko et al., 2014). Triaeris stenaspis has even been observed grasping springtail prey in a rodeo-riding manner (Korenko et al., 2014), as we observed in a juvenile P. pennimani capturing a large springtail.

Adult individuals of both sexes of P. pennimani and P. talquian continued feeding long after reaching maturity, with females even preying while guarding their egg sacs. For males, this continued feeding likely provides additional energy for mate searching. For females, it may have contributed to nutrients and energy reserves for oviposition and parental care. Some Lycosidae species feed during maternal care, albeit at lower rate, as noted by Nyffeler (2000). This is because it is well known that egg laying is an energy-consuming process (Foelix, 2011).

Retreat for molting. To our knowledge, no reports of molting retreat construction exist for any Phrurolithidae species. We therefore report this behavior in laboratory conditions for the first time, but this may also occur in natural conditions, although it is difficult to observe due to the small size of the specimens and their habit of living among leaf litter. This strategy is employed by both juvenile and penultimate individuals of P. pennimani and P. talquian, indicating that it is not restricted to early developmental stages. Individuals of both species appear to allocate resources to enhance their safety during the critical and vulnerable molting process. Not all spiders construct a retreat for molting; differences among families or even among species within the same genus have been documented (Austin, 1984a; Foelix, 2011; Nentwig & Heimer, 1987). Some Salticidae species, such as Servaea incana (Karsch, 1878) and Phidippus johnsoni (Peckham & Peckham, 1883), build a retreat for molting, although they also utilize it for courtship and as a refuge during breeding (McGinley et al., 2015; Jackson, 1979). The plasticity of the retreat constructed by both Phonotimpus species may be due to multiple factors, including the overall condition of the individuals and the structures available at their environment. For example, well-fed females of Tigrosa helluo (Walckenaer, 1837) are more likely to construct burrows than individuals with higher levels of hunger, as burrow construction can be an energy-intensive task (Walker et al., 1999).

Eggs sac, clutch size, and embryonic period. In spiders, the shape and conformation of an egg sac exhibit high variability, ranging from simple structures consisting of a few silk threads enveloping the eggs to elaborate egg sacs composed of multiple layers. In some cases, the egg sac is located within a retreat or nest (Foelix, 2011). Disk-shaped egg sacs appear to be common among phrurolithids, as evidenced by other genera such as Phrurotimpus sp. and an undescribed Phonotimpus species from central Mexico, which also construct similar egg sacs. Phrurotimpus sp. attaches its egg sacs to stones, while Phonotimpus secures them on dried leaves, with eggs clustered in the center (Kaston, 1948; unpublished data), resembling those of P. pennimani and P. talquian.

Although families related to phrurolithids produce egg sacs of diverse shapes, some exhibit similarities. For example, certain species within the genera Clubiona (Clubionidae), Castianeira Karsch, 1880 (Corinnidae), Agroeca Westring, 1861 (Liocranidae), Trachelas L. Koch, 1872 (Trachelidae), Callilepis Westring, 1874, Cesonia Simon, 1893, Herpyllus Hentz, 1832, Zelotes Gistel, 1848, and Drassodes Westring, 1851 (Gnaphosidae) also construct flattened oval or lenticular egg sacs (Kaston, 1948). Interestingly, all these related families inhabit microhabitats similar to those of phrurolithids.

The number of eggs laid, and the number of egg sacs constructed by a female spider vary among species, ranging from a single egg to thousands and from one to ten egg sacs (Kaston, 1948). Families related to Phrurolithidae, such as Clubionidae, Trachelidae, Corinnidae, and Gnaphosidae, produce between 8 and 60 eggs per egg sac (Kaston, 1948). The phrurolithids L. flavitarsis and P. festivus lay a mean of 4.7 and 4.1 eggs per clutch, respectively, and females of both species construct multiple egg sacs under laboratory conditions (Pekár & Jarab, 2011). In contrast, the Nearctic Phrurotimpus alarius produces 13 eggs per egg sac under laboratory conditions (Kaston, 1948; Montgomery, 1909). Available data indicate that phrurolithids, which are small-sized spiders, lay fewer eggs per egg sac but construct up to four egg sacs under laboratory conditions. Marshall and Gittleman (1994) reported a correlation between female spider body size and the number of eggs laid, with larger species producing more eggs. Finally, prior to this work, no data on the embryonic period for any phrurolithid species were known.

Maternal care. Parental care is any trait that enhances the fitness of a parent's offspring; for instance, by protecting them from predators, cannibalism, parasitoids, fungal infections, food shortages, and harsh environmental conditions (Santos et al., 2017; Smiseth et al., 2012; Tallamy, 1984). The most common forms of care in spiders involve egg sac construction, egg sac guarding, and brood care by the female (Yip & Rayor, 2014). Some spiders carry the egg sac or their offspring post-hatching (Kaston, 1948; Yip & Rayor, 2014), while others provide food, such as prey, regurgitated fluids, or trophic eggs, or engage in matriphagy (Ibarra-Núñez, 1985; Kim & Roland, 2000). However, most spiders typically abandon their egg sacs upon completion (Foelix, 2011).

Prior to Platnick's (2019) work, it was commonly accepted that phrurolithids abandoned their egg sacs. Specifically, Dondale and Redner (1982) stated that females of Phrurotimpus abandon their scale-like egg sacs. Based on the label of a female specimen collected by William Shear, Platnick (2019) asserted that Phrurotimpus borealis (Emerton, 1911) guards its egg sac and suggested the adoption of the common name "guardstone spiders" for all Phrurolithidae. Here, based on observations made in the field and under laboratory conditions, we found that females guard their egg sacs, and even when disturbed, they typically return to mount guard for most of the embryonic period. Additionally, field observations of an undescribed Phonotimpus species indicate that egg sac guarding might be shared by most, if not all, Phonotimpus. Some species related to phrurolithids, such as clubionids, gnaphosids, and salticids, also exhibit egg sac guarding until the emergence of offspring (Austin, 1984a; Kaston, 1948; Jackson, 1979).

The sole presence of a mother can increase the survival of progeny by deterring predators and parasitoids (Vieira & Romero, 2008). However, in some cases, such protection does not prevent parasitism (Barrantes & Weng, 2007) or is ineffective against larger predators like mantispids (McGinley et al., 2015; Vieira & Romero, 2008). Future studies should investigate the efficacy of guarding by females of P. pennimani and P. talquian in deterring natural enemies and reducing parasitism by Idris wasps or other natural enemies.

Endoparasitoids of P. pennimani . Here, we document the first record of an Idris sp. wasp (Scelionidae) parasitizing eggs of Phrurolithidae, with the wasp representing a putative new species of Idris (Margaría, 2019 in litt.). The wasp genera Baeus Haliday, Ceratobaeus Ashmead, Idris Förster, and Odontacolus Kieffer (Baeini, Scelioninae) specialize in exploiting spider eggs (Araneae) with a high degree of host specificity (Austin, 1984b; Austin et al., 2005; Barrantes & Weng, 2007; Loiácono & Margaría, 2013). Prior to hatching, the larva of scelinoid wasp consumes the contents of the host egg and pupates within it (Austin et al., 2005). In the Neotropical region, Idris spp. wasps are known to attack spider eggs of Ctenus sp. (Ctenidae), Pardosa ca. flavipalpis F. O. Pickard-Cambridge, 1902 (Lycosidae), and Sumampattus hudsoni Galiano, 1996 (Salticidae), among other unidentified spiders (Jiménez, 1987, Loiácono & Margaría, 2013).

The use of only two P. pennimani eggs by the Idris sp. wasp on each egg sac, may represent a strategy to avoid resource depletion. As observed in other Baeini (Scelioninae) wasps, the mortality of eggs caused by Ceratobaeus spp. and Baeus sp. wasps is limited to approximately 30% of total egg clutch in spiders such as Clubiona sp. (Clubionidae) and Theridion evexum Keyserling, 1884 (Theridiidae), respectively (Austin, 1984b, Barrantes & Weng, 2007). Austin (1984b) suggested that Baeini wasps can recognize their host using cues from the silk of egg sacs or retreats. Furthermore, it is known that Baeini female wasps can enter egg sacs to access eggs or oviposit by piercing the chorion through the thin silk of the host egg sac (Austin et al., 2005; Loiácono & Margaría, 2013). Future research should determine how the Idris sp. wasp locates Phrurolithidae egg sacs and whether Idris female wasps oviposit before or after P. pennimani seals her egg sac.

Acknowledgments.

G. M. Suárez (ECOSUR) provided vital assistance rearing spiders and springtails in laboratory. E. R. Chamé and G. M. Suárez (ECOSUR) help us with the acquisition of photos of live specimens and SEM images. E. R. Chamé and H. Montaño (ECOSUR) helped us in the field. A. Dor and L. Solís-Montero (ECOSUR) provided useful suggestions in a previous draft. J. G. Palacios Vargas (UNAM) determined all Collembola specimens. C. Margaría (Universidad Nacional de La Plata), M. Loiácono, and N. Johnson (The Ohio State University) corroborated the wasp identification. We thank G. Villegas, E. F. Campuzano and an anonymous reviewer for their suggestions and support during the review process. DCV was supported by a graduate scholarship from Consejo Nacional de Humanidades, Ciencias y Tecnologías (CONAHCYT).

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Notes

Author notes

*Corresponding author: Guillermo Ibarra-Núñez gibarra@ecosur.mx