Amphibians of North Carolina

Scientific Name:	Common Name:		Family (Alpha):

NC Records
Hylidae Members:
Hyla squirella - Squirrel Treefrog

Taxonomy

Class: Amphibia

Order: Anura

Family: Hylidae

Subfamily: Hylinae


Taxonomic Comments: Duellman et al. (2016) elected to split treefrogs in the genus Hyla into two genera. Hyla (sensu stricto) refers to a group of species that are found in Eurasia, while a new genus Dryophytes was resurrected to include all of the North and Central American species and three other species that are found in eastern temperate Asia. Members of these proposed genera cannot be distinguished by any known morphological features, but represent two clades. Whether members of these clades should be placed in separate genera or treated as lesser taxa such as subgenera is arbitrary and dependent on one's taxonomic philosophy. Here, we retain Hyla for North American species to be consistent with current usage by the Society for the Study of Amphibians and Reptiles.
Species Comments:

Identification

Description: Adult Hyla squirella are small treefrogs with short heads, smooth skin, and slender bodies. The dorsal ground color and patterning are highly variable, and individuals are often best identified by eliminating our other treefrog species. Individuals are often some shade of green, olive green, or yellowish-green, but are frequently darker shades such as gray, tan, brown, or dark brown. The dorsal ground color on the back and legs can vary from being unmarked to heavily marked with darker blotches. Blotching is generally most evident on brown specimens, and individuals can quickly change both the ground color and the prominence of the blotching on the dorsum. For example, a bright green individual with no spotting may change to brown with darker blotches in a matter of minutes (Wright 1932). Scattered yellow flecks are also sometimes present on the dorsum. An interocular dark bar is present on many specimens, and a white or pale line is often evident on each side of the head that extends along most of the upper lip. On many individuals this continues rearward across the shoulder, then as a diffuse, irregular streak towards the groin. When a streak is present, the lower edge is poorly defined. The venter is whitish, and there is often a yellowish wash in the groin and axilla region and in unexposed portions of the thighs. The males have a deep yellow throat in the breeding season, while the female throat is lighter. Males also have larger and brighter white stripes on the upper lip and sides (Buchanan 1994). The Squirrel Treefrog can be confused with other treefrogs, particularly the juveniles of our larger species. It superficially resembles the Green Treefrog, but has a diffuse whitish stripe along the side that fades into the ventral color below. In contrast, H. cinerea has a sharp, crisp stripe along the side that often has a thin, darker margin. The dark, blotched forms of H. squirella resemble H. femoralis, but H. squirella lacks the bold orangish-yellow spots on the rear of the thighs as seen on H. femoralis. This species also lacks the white blotch beneath the eye that is characteristic of H. chrysoscelis and H. versicolor. The adults vary from around 22-45 mm SUL, and the males in local populations usually average slightly smaller than the females (often < 5%, Dodd 2013). Some representative average SULs for males and females provided by Dodd (2013) include 33 mm and 33.4 (Charleston, South Carolina), 31.6 mm and 33.4 (Marion County, Florida), 33.9 mm and 31.5 (Miami, Florida), and 26.6 and 27.8 (Collier County, Florida). Males from Arkansas averaged 31.8 mm SUL (range = 29-35 mm) compared with 33.8 mm (range = 30-37 mm) for females (Connior et al. 2014). Beane et al. (2010) listed the size range of adults for the Carolinas and Virginia as 22-41 SUL, while Jensen et al. (2008) reported a size range of 22-45 mm for Georgia specimens. The tadpoles are drab colored, deep bodied, and reach a maximum size of 32–38 mm (Dodd 2013). A pale postorbital stripe is present and the tail is not distinctly striped. The tail fin and tail musculature are faintly marked with dark spots or blotches. The dorsal fin is lower in height than the tail musculature at any given point and terminates before reaching the region of the eyes. The tail tip is acuminate, and the throat and belly are yellowish with a black center (Dodd 2013).
Vocalizations: The males produce two common vocalizations -- the “rain call” or "tree call" that is made from trees during the day -- and the advertisement call that is made from the breeding sites at night. The advertisement call sounds like a nasally or raspy "rrraak" or "waaaak" and under field conditions is repeated in long, repetitive sequences at about two calls per second, "rrraak-rrraak-rrraak-rrraak". The rain call is somewhat similar and is best characterized as a half-hearted attempt at an advertisement call. The individual calls are issued at a slower rate (sounding like a raspy croak or quack of sorts) and often more irregularly when in a repetitive sequence. The repetitive sequences are usually of much shorter duration relative to the very prolonged repetitive advertisement calls. The rain call is given throughout the warmer months of the year, often with approaching rain or under conditions of high humidity. Its function remains a mystery. The advertisement call is used to attract mates and to advertise territories to other males. A male can produce over 6,000 advertisement calls per hour, and the energetic costs are very high (Dodd 2013, Prestwich et al. 1989). The shape of the vocal sac allows sound to be broadcast effectively in all directions.
Technical Reference: Dodd (2013)
Online Photos: Google iNaturalist
Observation Methods: The adults are commonly seen around the margins of wetlands during the breeding season, and can be found crossing roads on rainy nights. They are frequently seen around homes, buildings, and landscaped areas.

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AmphibiaWeb Account

Distribution in North Carolina

Distribution Comments: The Squirrel Treefrog is primarily confined to the Coastal Plain -- including numerous barrier islands -- from southeastern Virginia throughout all of peninsular Florida, then westward through Alabama, Mississippi, and Louisiana before ending in eastern Texas. The range extends northward in the Gulf region to extreme southeastern Oklahoma, extreme southern Arkansas, and through the Coosa Valley of central Alabama and an adjoining area of eastern Mississippi (Dodd 2013). Populations can be found outside of the Coastal Plain in the Carolinas and Georgia. In North Carolina, populations are common in the Coastal Plain and in some areas of the eastern and central Piedmont. This species appears to have expanded its range westward in the Piedmont in recent decades, as is the case in Georgia (Jensen et al. 2008), and is now well established in many areas of the eastern and central Piedmont. This and other treefrogs often hide in and on vehicles, boats, and nursery stock and can be transported long distances by humans. We have three records from the mountains that presumably reflect introductions from outside of the natural range.
Distribution Reference: Beane et al. (2010), Dodd (2013)
County Map: Clicking on a county returns the records for the species in that county.


GBIF Global Distribution

Key Habitat Requirements

Habitat: Except during the breeding season, the adults are commonly found in moist, forested habitats that are somewhat open. They have also adapted well to human-altered landscapes where breeding sites remain. Some of the commonly used habitats include deciduous or mixed-deciduous forests, upland oak forests, longleaf pine sandhills, mesic and xeric hammocks, coastal pine flatwoods, dome and basin swamps, wet prairies, and ecotones between forest and open areas (Dodd 2013). Individuals are commonly found around homes and other buildings where breeding sites such as flooded ditches occur nearby.

Breeding is mostly restricted to seasonal or semipermanent wetlands that lack fish, although small numbers of larvae have occasionally been found in habitats with fish. Sites that are sunny or only partially shaded are generally preferred over heavily-shaded sites (Binckley and Resetarits 2007). Natural habitats that are commonly used include small temporary pools, Carolina Bays, depression marshes, cypress savannas, and cypress-gum ponds (Babbitt et al. 2005, Dodd 2013, Liner et al. 2008). Populations have adapted well to human-disturbed landscapes and commonly use ditches and flooded agricultural fields. Populations are often found in coastal areas that receive salt spray, and the eggs and larvae seem to tolerate mildly brackish waters (Dodd 2013).

In North Carolina, the adults occupy a wide range of habitats. Natural habitats include Longleaf Pine savannas and flatwoods, maritime forests, bottomland forests, marshes, and pond edges. They can be quite common on barrier islands where they show some degree of tolerance for brackish waters and salt spray. Migrating individuals occasionally show up in old fields or wet pastures, and individuals are often found around outdoor lights at night where they feed on insects. Breeding takes place primarily in seasonal, fish-free habitats that range from small rain pools and mud puddles to large Carolina Bays. Breeding habitats include seasonal floodplain pools, shallow, ephemeral wetlands of various sorts, roadside ditches, seasonally flooded fields, borrow pits, and other fish-free habitats.

Environmental and Physiological Tolerances: Larvae are found in coastal areas affected by salt spray or occasional overwash and appear to tolerate mildly brackish water. Webb (1965) observed tadpoles in a shallow pool with a salinity of 4.7 parts per thousand in coastal North Carolina.

Adaptations to Natural Disturbances: Gunzburger et al. (2010) found that H. squirella did not recolonized wetlands that suffered overwash following a hurricane even though other native anurans did. This species appears to be well adapted to surviving in pinelands that are managed with prescribed burns (Langford et al. 2007).

Biotic Relationships: The larvae appear to lack chemical defenses against aquatic predators and are readily consumed by dragonfly larvae, giant water beetles, and fishes that occasionally colonize seasonal wetlands (Baber and Babbitt 2003, 2004, Babbitt and Tanner 1997, Binckley and Resetarits 2002). The tadpoles use chemical cues to monitor predators and respond by reducing activity levels (McCoy and Bowker 2008, Richardson 2001).

Adult choice of oviposition sites plays a critical role in reducing risks to a female's offspring. Binckley and Resetarits (2002, 2008) found that gravid females strongly avoided laying eggs in experimental pools with the Banded Sunfish (Enneacanthus obesus) that is a highly efficient predator on the tadpoles. The amount of nutrients that were added to pools did not affect the tendency of females to avoid fish pools, but they were more likely to use fish pools when large numbers of females were ovipositing on a given night.

The juveniles and adults are cryptically colored and can rapidly change their color and degree of dorsal blotching to blend in with ambient backgrounds (Dodd 2013). The predators of adults are poorly documented, but undoubtedly include aquatic and semiaquatic snakes, wading birds, raccoons and the like. Meshaka and Layne (2015) reported that this species is preyed upon by the the Peninsula Ribbon Snake, the Southern Leopard Frog, and the Cuban Treefrog. The latter can depress the sizes of local populations (Meshaka 2001).

See also Habitat Account for Coastal Plain Mixed Habitats

Life History and Autecology

Breeding and Courtship: The males begin calling following the spring warmup and typically continue through August or September. At southern locales they may continue calling sporadically into October or even later (Meshaka and Layne 2015). Populations may begin chorusing as early as February or March in southern populations (Brugger 1984, Carr 1940, Dodd 2013, Dundee and Rossman 1989) and at later dates farther north (Wright 1932). Calling in many populations does not begin in earnest until mid to late spring, and usually not until air temperatures exceed 19°C (Dodd 2013). Breeding in the Carolinas and Virginia typically occurs from April to August (Beane et al. 2010).

The arrival of warm seasonal rains triggers the adults to move to the breeding sites. They usually stay in the immediate vicinity of wetlands throughout the summer, then return to terrestrial habitats after breeding ends in late summer or early autumn (e.g., late September to November in Florida; Goin 1958, Boughton et al. 2000). Heavy, warm rains from thunderstorms or weather fronts usually trigger heavy breeding and the formation of large choruses (Wright 1932).

Much of our knowledge on aspects of courtship and mating is based on Brugger's (1984) detailed study in northern Florida that are summarized by Dodd (2013). In this region, the males arrive at breeding sites around sunset and most remain for only 2–4 hours. An entire chorusing event typically last 3–6 hours and most males only call for 1–2 hours per night. On most nights the females arrive 2-3 hours after sunset, but on very warm nights chorusing may not begin until after midnight. In such cases, the females arrive within 1–2 hours after males begin chorusing. As with most frogs, the operational sex ratios on a given night are highly biased towards males since they stay longer at the ponds. Brugger (1984) recorded ratios as high as 33:1 in Florida.

The males call from the ground surface on land and from clumps of grass, vegetation mats, and debris in shallow water near the shoreline. They also use vertical perches such as the stems of shrubs. The calling sites are generally no more than 1 meter above the water or land surface, or more than 1-2 meters from the edge of the pond when calling from land. The calling males stake out small territories and tend to be evenly spaced around a pond perimeter, with the distance between neighbors being around a meter or more. Calling males will issue an encounter (warning) call if an intruder approaches and the two may engage in a wrestling match to resolve contests (Dodd 2013). Non-calling, subordinate satellite males are sometimes present within or near a male's territory.

Females appear to select their mates based on call characteristics such as the call rate and pitch, and do not appear to select males based on their relative size. They may also use visual cues such as the degree of development of striping on the body (Taylor et al. 2007). A female will bypass satellite males as she moves towards a selected mate, and amplexus is initiated by her touching the calling male (Brugger 1984, Dodd 2013). Amplexus is axillary and the pair may remain amplexed for two or more hours before the eggs are laid and fertilized. Females typically take 35–65 minutes to lay their entire complement (Brugger 1984, Dodd 2013). Males may breed with more than one female during the breeding season (Brugger 1984), but females are thought to only breed once.

Reproductive Mode: Females lay their eggs singly in small groups that are scattered on the bottom or attached to grass blades (Brugger 1984, Wright 1932). Freshly laid eggs are brown above and white to cream below. Those that were examined by Wright (1932) averaged 0.9 mm in diameter (range = 0.8-1 .0 mm), with an inner envelop that averaged 1.4 mm (range = 1.4-1.6 mm) and an outer envelope that averaged 1.6 mm (range = 1.4-2.0 mm). The mean size of ova from three clutches from southern Florida averaged 1.76, 1.84, and 1.60 mm (Meshaka 2001). The embryos develop quickly, with hatching typically occurring within 2-4 days (Dodd 2013, Wright 1932).

A female rarely lays more than 2,000 eggs and typically far fewer. Two females from Georgia laid 942 and 972 eggs in captivity (Wright 1932). Eight females from Virginia (Mitchell and Pague 2014) laid an average of 1,076 eggs (range = 392–2,081), and four from South Carolina laid an average of 1,511 eggs (Hocking 2010). Brugger (1984) reported a mean of 1,059 eggs for five clutches observed in Florida, and a mean of 900 eggs per clutch (range 361–2,003) for specimens that were laid in captivity (Dodd 2013). Four specimens from Arkansas averaged 1,324 eggs (range = 701-1,635) based on ovarian egg counts (Connior et al. 2014), while three from south-central Florida had estimated clutch sizes of 808, 1,181, and 1,216 eggs (Meshaka 2001).

Aquatic Life History: Hyla squirella often breeds in warm, shallow seasonal pools that are subject to the vagaries of seasonal rainfall patterns. In some years the pools hold water until most or all of the tadpoles metamorphose, while in others the hydroperiods are relatively short and many or all are trapped and killed before they can metamorphose. The larvae have evolved several adaptations in response to these conditions. These include being active foragers, and having innately rapid growth rates and shorter larval periods compared to species that use permanent ponds as breeding sites. Rapid growth rates allow tadpoles to reduce predation from gape-limited predators and increase the likelihood of larvae escaping drying ponds.

The larvae often occur at relatively high densities in ponds and very likely compete for limited food resources (Babbitt and Tanner 1997). The ponds also have numerous predators such as odonate larvae. McCoy and Bowker (2008) found that the larvae lack inducible defenses as seen in some hylid frogs such as H. chrysoscelis. They also lack chemical defenses against predators and are palatable. Their primary defense is to reduce their relative activity levels in the presence of predators, which in turn reduces encounter rates (McCoy and Bowker 2008, Richardson 2001). The larvae become more active swimmers as they age, regardless of whether are exposed to predators or not. This likely reflects a tradeoff between the need to feed efficiently and grow rapidly, versus the need to reduce activity to avoid odonates and other predators. McCoy and Bowker (2008) also found that tadpoles that were previously exposed to predator chemicals were less likely to be killed compared to naive tadpoles. Structurally complex habitats with aquatic vegetation that provides cover also enhance larval survival (Babbitt and Tanner 1997).

The larval period of H. squirella is poorly documented but is thought to last around 1-2 months depending on site conditions, with tadpoles growing to as large as 32–38 mm TL before the initiation of metamorphosis (Dodd 2013, Meshaka and Layne 2015, Windes 2010, Wright 1932). Laboratory-reared tadpoles required an average of 21-31 days to metamorphose in an experiment conducted by Blouin (1992a), and had shorter larval periods than either H. gratiosa or H. cinerea. Larvae grown in field enclosures in Florida began metamorphosing after 28 days (Windes 2010). The mean SVL for 12 recently metamorphosed juveniles from Arkansas was 12.5 mm (Connior et al. 2014), while newly transformed froglets from Georgia varied from 11–13 mm SUL (Wright 1932). The smallest individuals that were observed by Meshaka (2001) in the Everglades were 12-14 mm SUL.

Terrestrial Life History: The young metamorphs begin moving away from the breeding ponds soon after they transform. Dispersal away from the ponds occurs at night and the young move to forested sites such as mesic hammocks or oak forests. They use retreats that are similar to those of the adults and most move into the tree canopy after completing their movements away from the breeding sites (Dodd 2013).

The young grow rapidly and many appear to reach sexual maturity within a year after metamorphosing. In the Everglades, metamorphs become sexually mature within a few months after metamorphosing when they reach a minimum length of 23 mm SUL (Meshaka 2001, Meshaka and Layne 2015). In southeastern Georgia, Wright (1932) concluded that individuals required 1-2 years to become sexual maturity, but this needs additional verification.

During the day the juveniles and adults rest and remain immobile with the limbs tucked close to the body and the eyes closed. They coat their bodies with a layer of mucous and lipids that retard water loss (Barbeau and Lillywhite 2005), and feed at night when diurnal predators are inactive and humidity is higher. Goin (1958) noted that they often rest on the undersides of oak leaves in the treetops. Individuals tend to move short distances from their daytime retreats to feed before returning to the same retreat used during the day. An individual may occasionally shift to a new site over an activity season. However, they generally tend to be highly philopatric to their home ranges, and will even return to a previously occupied site after overwintering elsewhere (Dodd 2013, Neill 1957a).

Although the diet is poorly documented, the juveniles and adults appear to be gape-limited, euryphagous predators that take a wide range of prey. Feeding occurs both when at the breeding sites and in the trees (Dodd 2013). Duellman and Schwartz (1958) found beetles, small crayfish, a spider, an ant, and a cricket in a few southern Florida specimens. In other specimens from southern Florida, flies and beetles were common prey -- but a wide range of other small invertebrates were eaten (Meshaka and Layne 2015).

Seasonal movements by the adults typically involve a yearly cycle of moving to the breeding sites after the spring warm-up, remaining in the vicinity of wetlands throughout the summer, then returning to terrestrial habitats by autumn (Dodd 2013, Goin 1958). Windes (2010) installed an array of pvc pipes around a wetland and found that the adults were highly philopatric. They were usually recaptured in the same pvc pipe after overwintering, which implies that individuals have small home ranges in woodlands that they return to after leaving to breed.

After breeding, the adults move into the surrounding forests and can be found considerable distances from water where they live in trees. Dodd (1996) found individuals in Florida from 446–914 m (mean = 594 m) from the nearest water body. Boughton et al. (2000) noted that they prefer deciduous trees over pines, and are usually found > 2 m off the ground. The rain calls are often issued from high in the forest canopy.

Frogs in Florida and other southern localities are active year-round (e.g., Zacharow et al. 2003), while those farther north are increasingly inactive depending on the severity of winter weather. During periods of cold or dry weather they seek refuge in protected sites such as tree cavities or under bark, logs, boards, and leaf litter. They will commonly use artificial refuges such as pvc pipes where they often reside in the same pipe for long periods of time (Campbell et al. 2010, Windes 2010, Wright 1932, Zacharow et al. 2003). Individuals generally become inactive when temperatures drop below 15°C (Dodd 2013). The arrival of warm seasonal rains during the spring triggers the adults to begin moving to the breeding sites (Boughton et al. 2000, Goin 1958).

General Ecology

Population Ecology: Local populations are centered around the breeding sites and may consist of a dozen or two adults at very small, isolated breeding sites to many hundreds of individuals where clusters of ponds are used locally (Dodd 2013). Brugger (1984) found that the adults frequently moved between local ponds, which suggests that populations using clusters of ponds may lack significant metapopulation structure at this scale. However, Windes (2010) provided evidence of strong site fidelity to a cluster of local ponds. It was uncertain if her study populations conformed best to a metapopulation model, or to a terrestrial habitat occupancy model.

Hether and Hoffman (2012) examined microsatellite variation in populations from Florida and southern Georgia and found that populations do show spatial structure. Two clusters were also evident that represented an eastern group and a second group in the Florida Panhandle. Models that included specific habitat types explained more variation in genetic distance than spatial distance alone, and upland oak habitats were important in providing connectivity between populations. The authors also found that mildly urbanization landscapes may increase gene flow between populations, perhaps because homes act as habitats for the juveniles and adults and there are numerous roadside ditches that serve as breeding sites.

Community Ecology: The Squirrel Treefrog often shares breeding ponds with many other anuran larvae, but ecological interactions between these species are poorly documented.

Adverse Environmental Impacts

Habitat Loss: Numerous populations of Hyla squirella have undoubtedly been lost since European colonization due to the destruction of natural wetlands and loss of native forests. These losses have been compensated for to some extent by the creation of artificial breeding sites such as flooded ditches.

Habitat Fragmentation: Hyla squirella is often found in open canopy forests and tolerates habitat fragmentation fairly well. This species is often found around homes and outbuildings and commonly occurs in urban and suburban environments. It can also be found regularly in agricultural landscapes where there are nearby forests and breeding sites such as flooded ditches. Despite being found in disturbed landscapes, populations have declined or are less frequently encountered in urban areas and areas with managed forests (Dodd 2013). Windes (2010) found that larvae perform better in ponds in seminatural pastures versus those in intensively managed pastures in Florida. Upland oak forests provide important habitats for the juveniles and adults.

Effects of Introduced Species/Induced Increases of Native Species: The Cuban Treefrog (Osteopilus septentrionalis) is an invasive species that was introduced to the Florida Keys and has now spread throughout all of peninsular Florida. This very large treefrog eats insects and native frogs, including H. squirella, and the tadpoles can impact native species by either outcompeting them for food resources or by consuming the eggs and hatchlings (Babbitt and Meshaka 2000, Meshaka 2001). Rice et al. (2011) showed that O. septentrionalis can depress populations of native frogs, including H. squirella.

Status in North Carolina

NHP State Rank: S5

Global Rank: G5

Environmental Threats: Urbanization and the continued loss of seasonal wetlands and forested habitats for the adults is the greatest concern for existing populations in North Carolina. This and other amphibians often cross roads on rainy nights and suffer significant mortality from vehicles.

Status Comments: The Squirrel Treefrog is widespread and often locally common in coastal regions of North Carolina, and appears to have expanded its range in the Piedmont. Populations show no evidence on widespread declines in the state, and may be on the increase in the Piedmont.

Stewardship: Local populations are best maintained by having clusters of seasonal ponds that hold water throughout the summer, along with suitable forest habitat for the juveniles and adults.

Recording Gallery for Hyla squirella - Squirrel Treefrog
	2022-05-25. Beaufort Co. Jim Petranka and Becky Elkin - An adult chorus with Gastrophryne also in the mix.

Photo Gallery for Hyla squirella - Squirrel Treefrog		40 photos are available. Only the most recent 30 are shown.
	Recorded by: B. Bockhahn Sampson Co.	Recorded by: John Petranka Wayne Co.
	Recorded by: Mark Basinger Wilson Co.	Recorded by: Mark Basinger Wilson Co.
	Recorded by: Travis McLain Cabarrus Co.	Recorded by: Mark Shields Onslow Co.
	Recorded by: K. Bischof Transylvania Co.	Recorded by: K. Bischof Transylvania Co.
	Recorded by: R. Newman Carteret Co.	Recorded by: S. Dagenhart Dare Co.
	Recorded by: Michael P. Morales Sampson Co.	Recorded by: Michael P. Morales Sampson Co.
	Recorded by: Michael P. Morales Sampson Co.	Recorded by: Jim Petranka, Becky Elkin, John Petranka, Sally Gewalt Beaufort Co.
	Recorded by: Jim Petranka, Becky Elkin, John Petranka, Sally Gewalt Beaufort Co.	Recorded by: Jim Petranka, Becky Elkin, John Petranka, Sally Gewalt Beaufort Co.
	Recorded by: Morgan Freese New Hanover Co.	Recorded by: Max Ramey Watauga Co.
	Recorded by: Morgan Freese Dare Co.	Recorded by: j.wyche Gates Co.
	Recorded by: J. Cox Dare Co.	Recorded by: j.wyche Gates Co.
	Recorded by: C. Bennett Dare Co.	Recorded by: K. Flood Dare Co.
	Recorded by: M. Gosselin, K. Flood Dare Co.	Recorded by: Jane Wyche Gates Co.
	Recorded by: Jane Wyche Gates Co.	Recorded by: K. Bischof Beaufort Co.
	Recorded by: K. Bischof Beaufort Co.	Recorded by: K. Bischof, M. Smith Beaufort Co.