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Female reptiles may be able to increase the fitness of offspring by selecting nest sites that are less vulnerable to human disturbances and predators, while providing an appropriate thermal and hydric environment for incubation of their eggs. Northern Pine Snakes (Pituophis melanoleucus) in the New Jersey Pinelands are the only snake in North America known to spend days digging their nests, exposing themselves to predators and people. Both nest site and the habitat selected are critical to ensuring reproductive success. Our overall objective was to determine the factors involved in nest-site selection by Northern Pine Snakes to determine trade-offs females face, and to determine whether these characteristics appear to have changed over the last 37 yr in response to human activities and habitat degradation. We compared characteristics of nest sites, random points within clearings, and random points in the surrounding forest for 42 Northern Pine Snake nests in 2020, and secondly, compared specific characteristics with those of 22 nests examined in 1983. The data indicate that (1) nest sites in 2020 were usually in soft sand, rather than in hard packed sand or in “sugar sand”; (2) sand penetration (e.g., sand density) of nests exhibited a narrow range, while random points were in sand ranging from sugar sand to hard packed sand; (3) the ground cover of nest sites was more often moss/lichen compared with other vegetation; and (4) nest sites were in more open habitats than random points within clearings or the surrounding forest. Compared to the 1980s, Northern Pine Snake nests examined in 2020 were in similar habitat conditions (intermediate sand type, open habitats, with surrounding forests), but were more often on paleodunes, deeper in forests, with higher rates of off-road-vehicle (ORV) disturbances. Females spend 2–4 d digging their nests, presumably to ensure appropriate conditions for offspring embryogenesis and to protect eggs and hatchlings. Our results indicate that, while digging, they are vulnerable to thermal stress, predation, poaching, and mortality from ORVs. Thus our results highlight the trade-off between the fitness of the nesting female and the fitness of her offspring.

Over a period of 4 mo, an entire collection of seven Pine Barrens treefrogs (Hyla andersonii) died or were euthanized after developing pallor, generalized edema, and coelomic effusion. Necropsy revealed large numbers of strongyloidid nematodes within the small intestines associated with a moderate mucosal hyperplasia. Strongyloides sp. parasitic females, representing a novel species, were isolated from the fixed intestinal tract. This case report represents the first full description of strongyloidiasis in a tree frog army and highlights the potential of Strongyloides spp. as a cause of rapid mortality events associated with protein-losing enteropathy in frogs.

Eastern Hognose Snakes (Heterodon platirhinos) are considered a species of conservation concern in the northeast United States because of their association with rare and declining habitats such as pine barrens and shrublands. These are disturbance-dependent habitats that currently require management to persist. We studied Eastern Hognose Snakes on a pitch pine–scrub oak barren in western Massachusetts from 2008 to 2013 to describe patterns of space use, habitat selection, and survival of this species and to evaluate the effects of habitat restoration and fuels management. We monitored 12 snakes with radio telemetry during the months of May to October 2008–2010. We examined habitat use versus availability using paired logistic regression analyses in which availability was temporally and spatially explicit in relation to radio-tracked snakes' previous use location and likely movements. We found that radio-tracked snakes significantly avoided closed-canopy forests and power line corridors, and inste...

The quality of habitat for a given species is fundamental to its persistence in that habitat space. Herbivorous insects often require a specific combination of host plants, floral resources, and physical features such as shelter. Identifying these different habitat features is a focus of ecology and conservation, particularly for managing rare or imperiled taxa. We investigated the patterns of microhabitat and host plant use of the rare frosted elfin butterfly, Callophrys irus, a larval host-plant specialist found in frequently disturbed sand plains, barrens, and sandhill pine-oak forests of the eastern United States. Previous studies have been conducted on populations in the Northeastern and Midwestern US, but the southern part of its range remains unstudied. Our efforts focused on a persistent C. irus colony in northeastern Florida, resulting in a geographically referenced census of larval host-plant Lupinus perennis, along with a multiple year survey of microhabitat features relevant to both C. irus adults and immatures. Results of the larval host-plant census revealed that the highest densities of host plants were located distant to the highest densities of C. irus. Hot-spot analysis confirmed the significance of this pattern, suggesting different habitat requirements for larval host-plant L. perennis and C. irus individuals in order to achieve maximum potential densities. Our survey of C. irus immatures showed a similar pattern of microhabitat affinity that was previously recorded in the Northeast and Midwestern US, including large larval host-plants, low amounts of ground cover vegetation, and the presence of some shade. Unique to our study we found that the presence of other herbivores of L. perennis such as larvae of the crambid moth Uresiphita reversalis had a negative effect on encountering C. irus immatures. Our results suggest that management that aims to conserve these species needs to include habitat factors that favor the overlap of these species and to consider where their densities are the highest.

Loss of wetlands throughout the southeastern United States threatens the persistence of the region's highly diverse freshwater fauna. Losses are especially concerning for rare species that maintain small or fragmented ranges, zoogeographies that characterize many of the region's numerous freshwater endemics. We assayed nuclear and mitochondrial DNA sequence data from the Pine Barrens Treefrog (Hyla andersonii), a rare species distributed across the Atlantic and Gulf coastal plains. We hypothesized that the species' evolutionary history has been associated with changes in wetlands during Quaternary interglacials and that the contemporaneous extent of wetlands is positively correlated with population genetic diversity. Genetic variation was highest in North Carolina and South Carolina and lowest in New Jersey and Florida. Mean times to common ancestry ranged from 132,486 to 1,290,605 yr before present, and effective sizes ranged from 4,241 individuals in New Jersey to 403,718 individuals in North Carolina. Population migration rates were generally very low (

I conducted anuran vocalization surveys in cranberry bogs, drainage ditches, and reservoirs of three active and three abandoned cranberry farms in the New Jersey Pinelands to examine the distribution of four anuran species in three consecutive years. I then applied a multinomial model to quantify their calling intensity and assess the influence of cranberry cultivation activities on each anuran species. I discovered that four species naturally inhabiting the Pinelands exhibited varied responses to cranberry cultivation. The responses ranged from positive to negative depending on the life history characteristics, disturbance tolerance, and especially the breeding season of the species in question. Green frogs (Lithobates (=Rana) clamitans) showed a high abundance in the bog habitats of active farms, where they exhibited prolonged courtship calls in late summer. Fowler's Toads (Anaxyrus (=Bufo) fowleri) exhibited high abundance in the ditches of active farms and the reservoirs of abandoned farms. O...

Timber Rattlesnakes, Crotalus horridus, are an endangered species in New Jersey, where high human population density places extraordinary pressures on natural populations. Disjunct populations are found in both the northern part of the state and a few scattered locations in the southern Pine Barrens. The Pine Barrens populations were previously shown to have reduced levels of genetic diversity compared to other populations in the Mid-Atlantic region. In this study, we used microsatellite DNA to examine the remaining six known C. horridus populations in the Pine Barrens and two in the northern part of the state. Bayesian analysis, genetic distance analysis, and factorial component analysis were performed, and the results indicated that the sampled populations represented four genetic subgroups, the two northern populations comprised one subgroup and the remaining six Pine Barrens populations comprised the other subgroups. Although there was evidence for isolation by distance among the Pine Barre...

Northern Pine Snakes (Pituophis melanoleucus) are one of the few snakes that spend the winter in underground hibernacula that they excavate. We report the use of hibernacula by Pine Snakes from 1986 to 2012 in the New Jersey Pine Barrens. We determined whether philopatry to a specific hibernaculum varied as a function of age, sex, and location of the hibernaculum. Three hibernacula were occupied nearly continuously for 27 yr by 1 to 27 snakes each year. With known-age snakes (N = 120), captured mainly as hatchlings and 2-yr-olds, we found that 23% were always philopatric. Philopatry was related to age of last capture, sex, and capture location. Philopatry was higher for 1) females compared with males, 2) snakes at two solitary hibernacula compared with a hibernaculum complex, and 3) snakes 6 yr old or younger, compared with older snakes. Of hatchlings found hibernating, 24% used the same hibernation site the next year, and 38% were located at year 4 or later. The number of snakes that always used the same hibernation site declined with the age of last capture. Snakes that entered hibernacula as hatchlings were found more often than those that entered as 2-yr-olds. For the seven snakes that were 14 yr or older, females were found 64- 86 % of the time, whereas males were found 15 to 50% of the time. Understanding the behavior and habitat requirements of snakes during different seasons is central to life-history analysis and for conserving viable populations.

We studied home range size and maximum dispersal distance from hibernacula in Northern Pinesnakes (Pituophis m. melanoleucus) at a 1418-ha preserve in Cumberland County, New Jersey, USA, between 1993 and 2003. We discovered 22 different winter hibernacula that were used by 39 Northern Pinesnakes. Of the 10 snakes monitored in hibernacula for 3–5 yr, shifting was observed by 8 individuals, and 2 females showed hibernacula philopatry for five consecutive years. The average minimum convex polygon home range of 14 radio-tracked Northern Pinesnakes was 105.51 ha (located 30–108 times/snake), whereas the average kernel density estimator home range was 50% isopleth = 38.99 ha and 90% isopleth = 133.15 ha. There were no differences in home range as a function of sex, but the number of years snakes were radio-tracked affected home range size. An adult male had the largest home range of 258 ha. The average distance traveled by radio-tracked Northern Pinesnakes from their winter hibernacula was 1321.0...

The decline in snake populations has been attributed primarily to anthropogenic disturbances, including habitat destruction and fragmentation. We radio-tracked Northern Pinesnakes (Pituophis m. melanoleucus) in a military Target Zone and in the surrounding undisturbed buffer zone at the Warren Grove Gunnery Range in the New Jersey Pine Barrens, USA, to examine differences in spatial movement patterns and behavior. There were no differences in home range size (X¯ = 60.0 ± 32.7 ha), core activity area (X¯ = 10.6 ± 8.2 ha), range length (X¯ = 1294.8 ± 32.7 m), or mean distance traveled per day (X¯ = 65.2 ± 30.1 m) across all snakes, or between sexes (male n = 13, female n = 11), or location within (n = 13) or outside (n = 11) of the Target Zone. However, there were differences between tracking years. Northern Pinesnakes in 2005 and 2006 (n = 13) traveled greater total distance and mean distance per day and had larger home ranges compared to snakes radio-tracked in 2003 and 2004 (...

Understanding the specific habitat requirements of reptiles during different life stages or seasons is critical to conserving viable populations. Northern Pinesnakes (Pituophis melanoleucus) are one of the few species that spend the winter in underground hibernacula, which they excavate themselves. We report on 26 years (1986–2011) of monitoring Pinesnake use at seven hibernacula in the New Jersey Pine Barrens. Our goal was to determine the frequency of repeated use, number of snakes present by year, disruptions of hibernacula, and the relationship between number of snakes present and the probability of occupancy of each hibernaculum in successive years. The overall goal was to determine the importance of protecting known hibernation sites regardless of whether they appear occupied in a given season. These data suggest that, if no snakes are observed entering a particular hibernaculum over a limited time period, it does not mean none are there or that none will use it in successive years. The var...

We monitored Common Nighthawk (Chordeiles minor) nests in a managed grassland in the New Jersey Pine Barrens in 2009 and 2010, and assessed habitat selection by comparing vegetation characteristics at nests with random locations. We found relatively high nest survival with an estimated 79% chance of survival through incubation (daily survival rate = 0.987, n = 16 nests); predation was the most common cause of failure (n = 2). Movements of young (up to 45 m from the original nest site) were frequent, which introduced uncertainty that prevented us from estimating survival through fledging. Nest sites had significantly more open ground cover (e.g., sand, lichen) than random sites, as well as less shrub and grass cover, shallower litter, and lower mean vegetation height.

We conducted mist-net surveys of migrating songbirds during fall migration 2007–2009 on the 1,300-ha Albany Pine Bush Preserve (APBP), a fire-managed inland pitch pine–scrub oak (Pinus rigida–Quercus spp.) barren in east-central New York. We banded 244 migrating passerines from 32 non-resident species in 8,610 net/m/hr documenting use of northeastern pine barrens as stopover sites for passerines with diverse breeding ecologies. We estimated the breeding site origin of six species (a kinglet, four warblers, and a sparrow) using stable hydrogen isotope measurements from flight feathers. There was a broad range of isotope ratios within each species indicating a large catchment area extending several hundred kilometers north and west of the stopover site. Over half the birds originated >750 km from the APBP. We found no evidence for geographical structure of the timing of migration through APBP; slopes of regression lines for capture date versus hydrogen isotope ratio from feathers (δDf) were not sta...

We studied a newly established population of Kirtland's Warbler (Dendroica kirtlandii) in Adams County, Wisconsin, nesting in a red pine (Pinus resinosa) plantation. We found eight males and five females in Adams County in 2008 and 10 males and 10 females in 2009. Five of seven (71%) males color-banded in 2008 returned in 2009, and at least eight successful nests produced an estimated 33 young over the 2 years. Red pine comprised 66.9% of trees on the main site, 20.6% were northern pin oak/black oak (Quercus ellipsoidalis/Q. velutina), and 12.5% were jack pine (Pinus banksiana). Total tree density at the main site was 1,876 trees/ha, lower than generally reported in Michigan. Percent canopy cover and ground cover types were similar to Michigan sites. Lowest live branch height of jack pine was similar to Michigan sites, but lowest live branches of red pine at our site were closer to the forest floor. Significant red pine die-off at our site combined with substantial natural jack pine recruitment c...

New York has 22 documented species of tiger beetles (Coleoptera: Cicindelidae). Over half of these species are considered rare, at risk, or potentially extirpated from the state. These rare species specialize on three sandy habitat types under threat from human disturbance: beaches, pine barrens, and riparian cobble bars. In 2005, we began a status assessment of eight of New York’s rarest tiger beetles, examining museum records, searching the literature, and conducting over 130 field surveys of historical and new locations. Significant findings included (1) no detections of four of the eight taxa; (2) no vehicle-free beach habitat suitable for reintroducing Cicindela dorsalis dorsalis; (3) C. hirticollis at only 4 of 26 historical locations; (4) C. patruela patruela at only one site statewide; and (5) C. ancocisconensis at only 3 of 28 de novo survey sites. Additional species that might be declining deserve our attention, as do some threats to tiger beetle habitats, like lack of beach wilderness, fire suppression in pine barrens, and river damming. Rarity in tiger beetles is a result of varying ecologies, which suggest different conservation strategies. Future inventory and documentation of tiger beetle occurrences need to take into account the metapopulation structure and imperfect detectability of these rare insects.

O presente trabalho teve como objetivo identificar os recursos vegetais do ecossistema restingautilizados na Restinga de Carapebus, Parque Nacional da Restinga de Jurubatiba, Rio de Janeiro, Brasil. Foram realizadas 13 visitas, com duração média de uma semana cada, na área de estudo e os informantes especialistas foram indicados pela população. Para cada espécie são apresentados o nome local, a categoria de uso, as indicações populares e o tipo de formação vegetal de sua ocorrência narestinga. Foi identificada apenas uma pessoa na região de Carapebus que conhece, utiliza e indicaas plantas da restinga para vários usos. O informante indicou 119 espécies utilizadas, distribuídas em 100 gêneros e 49 famílias, sendo as famílias Myrtaceae (10 espécies), Clusiaceae e Rubiaceae (ambas com 6 espécies) as mais representativas em número de espécies. As plantas utilizadas foram classificadas nas categorias: alimentar, medicinal, ornamental, tecnologia, higiênica, aromatizante,construção e combustível. As espécies de uso medicinal dominaram, com 45 representantes, entre as quais, a aplicabilidade antidiarreica contou com o maior número de espécies (10). A maioria das espécies utilizáveis (67) ocorre na formação arbustiva aberta de Clusia. Os resultados apresentados neste trabalho são de fundamental importância para a preservação cultural da utilização dos recursos vegetais na Restinga de Carapebus.

In light of the recent discovery of an unrecognized species within nominal Zale lunifera (Hubner), the taxonomy of Z. lunifera is reassessed. Zale intenta (Walker), stat. rev. is the name that applies to the widespread species previously called Z. lunifera. Zale lunifera sensu stricto is the species previously thought to be undescribed; it occurs from the southern Atlantic coastal plain northward to the pine barrens of north-eastern United States. A differential diagnosis and adult illustrations of the two species are given.

Centrarchids (sunfish and bass species) and Bullfrogs (Lithobates catesbeianus) have been introduced into aquatic systems around the world and have the potential to negatively impact native fish and anurans. We surveyed fish and anuran assemblages from 26 impoundments in the New Jersey Pinelands, USA. We excluded non-native species and used ordination analysis to generate four native-species community gradients based on native-fish and anuran presence–absence and abundance data. All four community gradients paralleled an increase in the percentage of upstream-altered land (development and upland agriculture) and an increase in either non-native-centrarchid richness or abundance. Based on presence–absence and abundance data, native-fish and native-anuran assemblages, including restricted species (those generally limited to the Pinelands region) and widespread species (those widely distributed in New Jersey), differed between impoundments with and without non-native centrarchids. Compared to widesp...

A previously unknown species of the North American leafhopper genus Flexamia, Flexamia whitcombi sp. n., is described from pinebarren smokegrass (Muhlenbergia torreyana (Schult.) Hitchc.), a state-listed threatened grass in the New Jersey Pine Barrens. The serrata species group, to which it belongs, is redefined and a key to the species of the group is provided. This is the first reported insect association with Muhlenbergia torreyana.

One hundred and thirty-three species of vertebrates were recorded from areas representing three different landsystems on a grazing property on the Paroo River, located 52 kilometres south of Wanaaring in the Western Division of New South Wales. Fauna diversity was highest along the riparian zone of the Paroo River (the Paroo landsystem) and lowest on the red sand plains of the Gumbalara landsystem. The Cobham landsystem, of which the wetland environment of Waitche Lake is an example, gave an intermediate level of species diversity. The Paroo landsystem had the greatest number of small birds, which may have been indicative of the presence of a dense understorey and shrub layer in conjunction with a distinct tree canopy. Overall waterbird abundance and diversity was low due to flood conditions and local rainfall providing ample feeding and nesting areas, thus spreading populations over a greater area. Three threatened species (Brolga. Pink Cockatoo, Yellow-bellied Sheathtail Bat) were recorded as well as a ...

We predicted that bird diversity and abundance of some bird species would be higher in old-growth stands than in mature pine stands because of the greater structural diversity in old growth. We also predicted that patch size of stands should be influential. To test these predictions, we modeled counts of 79 bird species from 52 stands in 5 regions in the province of Ontario, Canada in relation to habitat at the local and landscape extents. Neither total species richness nor abundance differed between stand types. No significant difference was found in bird assemblages between stand types using ordination analysis. However, more Neotropical migrants were found in old-growth stands than in mature stands, while the reverse was true for short-distance migrants. Twenty-five species had higher counts in old-growth stands—three significantly so: Brown Creeper Certhia americana, Northern Parula Setophaga americana, and Scarlet Tanager Piranga olivacea. Supercanopy pine (> 60 cm dbh) was a significant (P 40 cm/dbh) was a significant positive predictor for Brown Creeper, Pine Warbler Setophaga pinus, and total species richness. The density of supercanopy and medium/large pine explained a small but significant amount of variation in bird assemblages (1%), after considering age, other tree variables (9%), and landscape metrics. Patch size was significant for Evening Grosbeak Coccothraustes vespertinus and total abundance. According to receiver operating characteristic (ROC) thresholds, Brown Creeper required a minimum of 62 stems/ha of medium/large pine. Pileated Woodpecker Dryocopus pileatus and Black-throated Green Warbler required a minimum of 14 and 23 stems/ha of supercanopy pine, respectively. Blackburnian Warbler Setophaga fusca required a minimum stand age of 66 years. Current targets in shelterwood seed cuts for pine appear to be just within range for Brown Creeper—at least for the first cut, but not for subsequent cuts. We recommend that forest management seek to emulate increased old-growth characteristics in a proportion of managed stands that fall within the range of variation expected under natural variation. RESUME. Nous avons prevu que la diversite et l’abondance de certaines especes d’oiseaux seraient plus grandes dans les vieux peuplements de pins comparativement aux peuplements matures a cause de la plus grande diversite structurale des premiers. Nous avons egalement prevu que la superficie des peuplements aurait une influence. Pour verifier ces hypotheses, nous avons modelise les denombrements de 79 especes d’oiseaux dans 52 peuplements de 5 regions de l’Ontario, au Canada, en relation avec l’habitat a l’echelle locale et a celle du paysage. Ni la richesse specifique totale ni l’abondance n’ont differe entre les types de peuplements. L’assemblage aviaire n’etait pas significativement different entre les types de peuplements, selon une analyse par ordination. Toutefois, un nombre superieur de migrateurs neotropicaux a ete trouve dans les vieux peuplements, tandis que les migrateurs de courte distance l’emportaient dans les peuplements matures. Vingt-cinq especes etaient plus abondantes dans les vieux peuplements, dont trois de facon significative : le Grimpereau brun Certhia americana, la Paruline a collier Setophaga americana et le Piranga ecarlate Piranga olivacea. Les pins de tres grande taille (> 60 cm dhp) representaient une variable explicative positive significative (p 40 cm dhp) representaient une variable explicative positive significative pour le Grimpereau brun, la Paruline des pins Setophaga pinus et la richesse specifique totale. La densite des pins de tres grande taille et celle des pins de moyenne ou grande taille ont explique une faible proportion – mais significative – de la variabilite de l’assemblage aviaire (1 %), apres qu’aient ete consideres l’âge, d’autres caracteristiques liees aux arbres (9 %) et des parametres du paysage. La superficie des peuplements etait significative pour le Gros-bec errant Coccothraustes vespertinus Aquila Conservation and Environment Consulting, Canadian Forest Service, Ontario Ministry of Natural Resources Avian Conservation and Ecology 7(1): 4 http://www.ace-eco.org/vol7/iss1/art4/ et l’abondance totale. D’apres les seuils ROC, le Grimpereau brun avait besoin d’un minimum de 62 tiges/ha de pins de moyenne ou grande taille, le Grand Pic Dryocopus pileatus et la Paruline a gorge noire recherchaient un minimum de 14 et 23 tiges/ha de pins de tres grande taille, respectivement, et la Paruline a gorge orangee Setophaga fusca avait besoin de peuplements âges d’au moins 66 ans. Les prescriptions actuelles des coupes progressives d’ensemencement pour les pins semblent etre tout juste suffisantes pour le Grimpereau brun – pour ce qui est de la premiere coupe, mais pas pour les coupes subsequentes. Nous recommandons donc que l’amenagement forestier cherche a reproduire les caracteristiques des vieilles forets dans une proportion de peuplements amenages qui correspond aux variations attendues dans les conditions de variations naturelles.

We studied nesting behavior of pine snakes Pituophis melanoleucus from 1977 to 1989. Female pine snakes in the New Jersey Pine Barrens excavate tunnels and a nest chamber for egg deposition. Over 70% of all nests had old, hatched shells remaining in the nests from previous years. The presence of hatched shells and the incidence of recaptured females in the same nest indicates nest site tenacity for some females. From 1977 to 1988 we branded 98 females and 45 were recaptured in the same general nesting area. All 45 recaptured females were recaptured one to four times at the same nest site; two of these snakes were captured in three years at the same site and in one year at a different site. We conclude that females show a high degree of nest site philopatry. We suggest that high nest site philopatry is selected for because it promotes location of sites that are safe from predators (buried, hatched shells) and contain suitable temperature conditions.

In 1993, we conducted an anuran-vocalization survey of 87 sites within two river systems (Mullica River and Wading River) of a major Coastal Plain watershed in the New Jersey Pinelands. The Mullica River system is more intensely developed and farmed than the Wading River system. Degraded waters, characterized by elevated pH, specific conductance, and nitrate-nitrogen values, are more frequently encountered in the Mullica River system. We found differences in the distribution of both individual anuran species and species-based site classes between the two river systems. Six native-Pinelands species, the Pine Barrens treefrog (Hyla andersonii), the carpenter frog (Rana virgatipes), the southern leopard frog (Rana utricularia), Fowler's toad (Bufo woodhousii fowleri), the spring peeper (Pseudacris crucifer), and the green frog (Rana clamitans melanota), were widely distributed, although spring peepers occurred more frequently in the Mullica River system. Four species that are distributed outside the region, but do not usually occur in the Pinelands, were heard only in the Mullica River system. These non-native Pinelands species were bullfrogs (Rana catesbeiana), pickerel frogs (Rana palustris), northern cricket frogs (Acris crepitans crepitans), and gray treefrogs (Hyla versicolor). Bullfrogs occurred more frequently at sites with adjacent developed or upland-agricultural land, while carpenter frogs were more closely associated with unaltered sites. Pine Barrens treefrogs and carpenter frogs were generally absent and leopard frogs occurred infrequently at sites where bullfrogs occurred. al of Herpetology, Vol. 34, No. 2, pp. 210-221, 2000 i t 20 0 Society for the Study of Amphibians and Reptiles istribution of Anurans in Two River Systems of a Coastal Anthropogenic factors have contributed to declines in amphibian populations in many areas (Wake and Morowitz, 1991). Causal hypotheses for local or regional declines in amphibian Corresponding author. species include wetland habitat loss (Lannoo et al., 1994; Hecnar, 1997), landscape fragmentation (Laan and Verboom, 1990; Gulve, 1994), aquatic pollution (Berger, 1989; Baker and Waights, 1993, 1994; Hecnar, 1995; Materna et al., 1995; Berrill et al., 1997), the presence of non210 This content downloaded from 157.55.39.217 on Mon, 05 Sep 2016 05:00:20 UTC All use subject to http://about.jstor.org/terms COASTAL PLAIN ANURANS native amphibians or predatory fish (Hammerson, 1982; Hayes and Jennings, 1986; Sexton and Philips, 1986; Bradford, 1989; Hecnar, 1997), and disease related to environmental stress (Carey, 1993; Crawshaw, 1997). The association between anthropogenic stresses and amphibian declines suggests that amphibians may be good indicators of environmental conditions (Barinaga, 1990; Wake, 1991; Wake and Morowitz, 1991; Blaustein, 1994; Hecnar and M'Closkey, 1996a), although the general applicability of this assumption has been disputed (Pechmann and Wilbur, 1994). The amphibian fauna of the Pinelands (Pine Barrens), located in southern New Jersey, is distinct from other areas of the state (Gosner and Black, 1957; Conant, 1979). In New Jersey, Pine Barrens treefrogs (Hyla andersonii) and carpenter frogs (Rana virgatipes) are found only in Pinelands habitats. Other Pinelands species, such as southern leopard frogs (Rana utricularia), Fowler's toads (Bufo zwodhousii fowleri), spring peepers (Pseudacris crucifer), and green frogs (Rana clamitans melanota), are also found throughout the southern portion of the state. Anuran species that are widely distributed outside the region, but do not usually occur in the Pinelands, include bullfrogs (Rana catesbeiana), pickerel frogs (Rana palustris), northern cricket frogs (Acris crepitans crepitans), and gray treefrogs (Hyla versicolor). These species may be found in Pinelands habitats that have been disturbed by humans (Gosner and Black, 1957; Conant, 1979). The general absence of these non-indigenous species in undisturbed areas of the Pinelands has been attributed to the high acidity of surface waters, among other factors (Gosner and Black, 1957; Freda and Dunson, 1986). The presence of bullfrogs in the Pinelands may be especially significant because predation or competition by this species has been implicated in the decline of native amphibian populations in other parts of the United States (Moyle, 1973; Bury and Luckenbach, 1976; Hammerson, 1982; Schwalbe and Rosen, 1988; Kupferberg, 1997). Assessing the status of amphibians in relation to anthropogenic factors requires a regional viewpoint (Hecnar and M'Closkey, 1996a). In this paper, we describe the results of an anuran survey conducted in 1993 within two river systems (Mullica River and Wading River) of a major Coastal Plain watershed located entirely within the Pinelands. We address four major questions. First, are there differences in the distribution and abundance of individual Pinelands and non-Pinelands anuran species between the two river systems? Second, is the distribution and abundance of individual anuran species related to land-use disturbance? Third, what is the association between the bullfrog and Pinelands anuran species? Fourth, are there differences in the species composition of sites between the two river systems? MATERIALS AND METHODS Study Area.-We conducted our survey in the 1473-km2 Mullica River basin (Fig. 1). This watershed comprises about one-third of the Pinelands, is centrally located in the region, and displays a range of land-use characteristics, including forest and highly developed and farmed areas. It lies entirely within the Pinelands on the Outer Coastal Plain and is associated with the Kirkwood-Cohansey aquifer (Rhodehamel, 1979; Johnson and Watt, 1996). The Mullica River b sin includes two major river systems, the Mullica River and the Wading River (Fig. 1). These two river systems display distinctly different land-use patterns and associated waterq ality characteristics. The Mullica River system, located on the western side of the Mullica River basin, is more intensely developed and farmed than the Wading River system, located on the eastern side of the basin (Fig. 1). Most alt r d land in the Wading River system is farmed wetlands. Differences in land-use patterns are reflected in stream water quality (Fig. 1). Stream waters in the least disturbed Pinelands drainage basins are usually acid and low in dissolved solids compared to waters affected by development and upland agriculture (Morgan and Good, 1988; Zampella, 1994; Johnson and Watt, 1996). Degraded waters, characterized by el vated pH, specific conductance, and nitrate-nitrogen values, are more frequently encountered in the Mullica River system (Fig. 1). Non-indigenous plant and fish species also occur in degraded streams in this portion of the Mullica River basin (Hastings, 1984; Zampella and Laidig, 1997; Zampella and Bunnell, 1998). Study Sites.-We surveyed 39 sites in the Wading River system and 48 sites in the Mullica River system (Fig. 2). Survey sites included three forested and 26 non-forested stream sites, 27 stream impoundments, six abandoned bogs, 14 borrow pits (excavated ponds), and 11 ponds found in naturally occurring depressions (Table 1). All stream sites were located at road crossings. Forested stream sites were bordered by red maple (Acer rubrum) or Atlantic white cedar (Chamaecyparis thyoides) swamps. Non-forested stream sites included small impoundments ( 0.5 ha), open water habitats. The abandoned-bog group included old cranberry bogs or impoundments that have succeeded to mixed shrub /emergent wetlands with open water. All abandoned bogs were historically connected to streams. The borrow pits 211 This content downloaded from 157.55.39.217 on Mon, 05 Sep 2016 05:00:20 UTC All use subject to http://about.jstor.org/terms R. A. ZAMPELLA AND J. F. BUNNELL Mullica River System Specific conductance (gS/cm)

iables in 14 acid water ponds located within drainage areas that displayed a range of agricultural and developed land use cover in the New Jersey Pinelands (Pine Barrens). Species were classified as those restricted to the Pinelands (Pine Barrens species: Hyla andersonii and Rana virgatipes), those with a wide-ranging distribution in southern New Jersey (wide-ranging species: Pseudacris crucifer crucifer, Bufofowleri, Rana sphenocephala utricularius, and Rana clamitans melanota), and those normally unable to enter the Pinelands except in habitats disturbed by human activities (border-entrant species: Hyla versicolor, Pseudacris kalmi, Acris crepitans crepitans, and Rana sylvatica). The frequency of occurrence of Pine Barrens and wide-ranging species was greater than that of any border-entrant species. Detrended correspondence analysis revealed an adult community gradient associated with geographic position where border-entrant species were heard only at ponds located along the transition between forested and disturbed landscapes. Principal component analysis revealed a primary environmental gradient of increasing pH and emergent herbaceous vegetation cover and decreasing specific conductance and floating vegetation cover and contrasted natural and artificial (borrow pits) pond types. Floating vegetation cover and specific conductance were lower and pH was higher in artificial versus natural ponds. Larval recruitment occurred for all species except the four border entrants and B. fowleri. Larval species richness was directly related to emergent herbaceous vegetation cover and inversely related to specific conductance. Our results suggest that landscape patterns influence the distribution of adult anurans, whereas pond chemistry may limit recruitment.

FORD, N. B., AND R. A. SEIGEL. 1989. Phenotypic plasticity in reproductive traits: evident from a viviparous snake. Ecology 70:1768-1774. LUTTERSCHMIDT, W. I., AND H. K. REINERT. 1990. The effect of ingested transmitters upon the temperature preference of the northern water snake, Nerodia s. sipedon. Herpetologica 46:39-42. REINERT, H. K. 1984. Habitat separation between sympatric snake populations. Ecology 65:478-486. , AND D. CUNDALL. 1982. An improved surgical implantation method for radio-tracking snakes. Copeia 1982:702-705. , AND R. T. ZAPPALORTI. 1988. Timber rattlesnakes (Crotalus horridus) of the Pine Barrens: their movement patterns and habitat preference. Copeia 1988:964-978.

We studied the excavation and nesting behavior of pine snakes (Pituophis m. melanoleucus) in the New Jersey Pine Barrens from 1977-1989. Females excavate a tunnel and nest chamber for egg laying. Females initially make slight depressions in the sand (pre-test holes, 22%), then construct small test holes (16%), and finally excavate long tunnels leading to nest burrows (62%). Test holes and nests did not differ in most external characteristics (dump pile size, entrance size), although they did differ in soil moisture at the end of the tunnel, mean tunnel length, and number of turns. Mean clutch size was 9.5 ? 0.3 (range 4-16), and was related to female snout-vent length (r = 0.60, P < 0.001). Female vertebrates exert control over the growth, development, and survival of embryos either by having internal development (i.e., most mammals), impregnable eggs (i.e., most birds)

Although Central America has been investicount of ecological conditions in the pine sagated by biologists and worked by professional vanna and the adjacent broad-leafed forest and collectors for over a century, the region inh ave discussed species diversity, population cludes a large area of distinctive habitat that density, intraand interspecific competition, has remained almost unknown zoologically and niche utilization in the birds of these conuntil recent years. This pine savanna includes trasting habitats. This paper deals primarily only a single species of pine, Pinus caribaea; with the taxonomy, distribution, and reproducthe region resembles the pinelands of British tive biology of those species that use the disHonduras, but is entirely of low relief-gentinctive features of the pine savanna in some erally less than 100 m above sea level. The ecologically important way-i.e., foraging, Mosquitia is effectively isolated from the feeding, resting, nesting-with at least some British Honduras pinelands and from the regularity. I have therefore not listed every highland pine forests of Honduras and northspecies recorded in or over the savanna, but central Nicaragua by wide expanses of humid, have included some that may not use the salowland, tropical, broad-leafed forest. The vanna regularly or importantly but which are approximate extent of the savanna is shown in of distributional or taxonomic interest. figure 1; the southern extent of this habitat, beThose species using the savanna importantly tween Laguna de Perlas and Bluefields, Nicmay be categorized as follows: aragua, marks the southern limit of naturally Permanent residents-resident species whose occurring pines in the Western Hemisphere. presence depends on the presence of the saThe paucity of published biological informavanna, which carry on most or all of their tion on the Mosquitia until the past few years activities there, and which are absent from is striking. The few visits made by ornitholoadjacent areas of broad-leafed forest. gists to the Nicaraguan area up to 1963 have Wide-ranging residents-resident species been discussed by Howell ( 1965). Studies that use the savamra frequently and imporby Parsons ( 1955)) Radley ( 1960)) Johannestantly, perhaps exclusively in the case of some sen ( 1963), Taylor (1963), and Munro ( 1966) individuals, but which (as species) also use provide valuable information on the geography adjacent nonsavanna habitats frequently and an d ecology of the region; the most recent importantly. zoological studies dealing with the pine saSummer residents-differ from permanent vanna are those of Monroe (1968) in Honresidents only by their absence during the duras, Howell ( 1965), Campbell and Howell period from November to March, at least. ( 1965)) Monroe and Howell ( 1966), and Winter residents-species that nest in temBuchanan and Howell (1965). perate North America and are found regularly From 1963 to 1967, my associates and I in the savanna during the winter months. made five visits to the Nicaraguan savanna Regular visitors from adjacent broad-leafed region, as follows: 16 January to 1 February forest-resident species that are widespread 1963; 11 to 27 August 1965; 10 to 26 March in broad-leafed forest or forest edge, but 1966; 18 November to 1 December 1966; and which regularly range into the savanna and 15 to 30 April 1967. The personnel included 0. may even nest in the pines; however, all these M. Buchanan, L. F. Kiff, M. B. Lloyd, F. G. species depend on the adjacent broad-leafed Stiles, and J, E. Zoeger. These visits included forest for many or most of their activities. periods in both the wet and dry seasons, pePassage migrants-spring or fall transients riods when some spring and fall transients and that pass through the savanna. winter residents were present, and periods These categories may be abbreviated as PR, when breeding birds that are not year-round WrR, SR, WR, RV, and M, respectively, and residents were present. In another paper these abbreviations are used where applicable (Howell 1971), I have given a detailed acin the species accounts. Equivocal cases and

Two natural hybrids of Hyla andersonii are reported from Okaloosa County, Florida. Chromosome analyses confirmed the hybrid nature of the specimens and identified the parent species. The chromosome complement of each hybrid shows the presence of nucleolar-organizer regions unique to each of the two respective parents. Meiotic observations indicate that both hybrids were infertile. These hybrids cross previously proposed species groups of Hyla based on artificial hybridization experiments.

Highlight: Potential forage production is higher in the South than in other range areas of the United States, although actual production is declining rapidly due to accelerated pine regeneration. The cutover longleaf (Pinus palustris Mill.) pinelands that produced an abundance of forage have been largely regenerated with fast-growing slash (P. elliottii Engelm.) and loblolly pines (P. taeda L.) and these young plantations reduce herbage production drastically within a few years. Few large industrial timber companies encourage grazing, although some allow it, often without fee, as a public relations gesture. Cattlemen who depend on forest range alone seldom own the land their cattle graze, often lease the land under an annual permit, and have little incentive to improve the range. Attempts to promote cooperation among livestock producers through grazing associations have generally been unsuccessful. Public land managers are under pressure from wildlife and environmental organizations to prohibit or curtail grazing. Operational-scale multiple-use research is needed to evaluate compatibility of cattle, wildlife, and other resources.

We examined nest site selection by 31 female pine snakes that excavated and laid eggs at 22 nest sites in the Pine Barrens of southern New Jersey. All nests occurred in large clearings with less than 10% tree cover in pitch pine-scrub oak uplands. Nest sites differed from randomly selected points adjacent to nests with respect to vegetation cover and composition, distance to clearing edges and soil characteristics. Nests were in soft sand in open, unvegetated sections of clearings. There were few differences between solitary and communal nests except that more solitary nests were in Pennsylvania sedge than in grass in areas with greater tree cover and closer to roads.

Selected naturally regenerated flatwoods forests were burned in preparing a large, long-term study of the effects of several multiple use management practices on forest vegetation and wildlife. Early effects of burning on understory vegetation are reported here. Fire reduced woody understory coverage (from 72 to 66% of surface area), and increased herbaceous species frequency (from 60 to 81%) and herbaceous standing biomass (from 124 to 245 kg/ha). Graphical analyses show an increase in herbaceous species diversity as a result of burning. Successful production of wood, cattle, and wildlife in the southern pinelands depends on methods of managing both overstory and understory simultaneously to increase multiple product yields while avoiding environmental and ecological degradation. In 1976 a large study was begun to examine various effects of an array of forest management practices on wildlife, wildlife habitat, and herbage production. Portions of the study areas have only been burned. Reported here is an early assessment of the effects fire produced on the understory vegetation. Our objective is to show in a simple but compelling way certain beneficial effects of prescribed fire in a forest ecosystem. Study Area and Methods The 73-ha experimental area is on the University of Florida's Austin Cary Memorial Forest near Gainesville, Alachua County, Fla. When this study began the area was occupied by a 50-year old mixed stand of slash and longleaf pine averaging 20.7 m2 of basal area per ha. (This overstory was apparently not altered by the fires described in this paper.) Soils on about 70% of the experimental area are mainly a sandy, siliceous, hyperthermic Ultic haplaquod of the Pomona series. These soils are acid sands that contain an organic stain layer between 41 and 61 cm, and a clay layer at about 109 cm. Basinger fine sand (a siliceous, hyperthermic Spodicpsammaquent), which occurs at slightly lower elevations, occupies about 15%. Sparr sand (a loamy, siliceous, hyperthermic Grossarenic paleudult) and Adamsville sand (a hyperthermic, uncoated Aquic quartzipsamment) occur at slightly higher elevations and occupy most of the remaining 15% of the area. The entire experimental area was prescribe-burned during the winter of 1975-76. Prior to that, the forest had been protected from fire and other disturbances for many years, and consequently the understory was dominated by shrubs, mainly saw-palmetto and common gallberry. Bracken fern was plentiful and pineland threeawn was common in open spaces. Other herbaceous species were scarce and hard to find. Burning was designed to consume as much surface litter and understory biomass as possible with minimum damage to overstory vegetation. The objectives were to improve light penetrations and to expose seeds of desirable plants to mineral soil. Burning took place during February, two or three days after a rain while the Authors are research wildlife biologist (retired) and research forester, Southeastern Forest Exp. Sta., U.S. Forest Serv. and range research biologist, School of Forest Resources and Conservation, Univ. of Florida, Gainesville 32611. soil was moist, air temperature cool, relative humidity high, and wind light and steady. The results were extensive dieback of many shrubs, but shrub consumption by flames was far from complete. In September and October of 1976, six 1-ha plots randomly dispersed over the experimental area were selected for subsequent burning and the understory vegetation on these plots was surveyed. Three 30-m line transects were randomly located in each plot and monumented so they could be relocated. Coverage of all woody vegetation less than 1.5 m aboveground was determined by measuring to the nearest centimeter the portion of each transect covered by crowns of each woody species. In those spaces where woody plants did not cover the transect, measureTable 1. Canopy coverage (%) below 1.5 m of woody plants on line transects following prescribed fires in February 1976 and in February 1978.

AND . 1988. Habitat use in free ranging pine snakes Pituophis melanoleucus in New Jersey Pine Barrens. Herpetologica. 44:48-55. FORMAN, R. T. T. 1979. Pine barrens: ecosystem and landscape. Academic Press, New York. GIBBONS, J. W., AND R. D. SEMLITSCH. 1987. Activity patterns. In R. A. Seigel, J. T. Collins, and S. S. Novak (eds.), Snakes: ecology and evolutionary biology, pp. 396-421. Macmillan, New York. GREGORY, P. T. 1983. Identification of sex of small snakes in the field. Herp. Rev. 14:42-43. HEATWOLE, H. 1977. Habitat selection in reptiles. In C. Gans and D. Tinkle (eds.), Biology of the Reptilia, Vol. 7, pp. 137-155. Academic Press, New York.

We studied sexual and individual variation in habitat use by pine snakes, Pituophis melanoleucus (5 males, 5 females), in the New Jersey Pine Barrens. Both sexes were found in pine-oak forests. Habitat selection varied by sex and individual, with males using logs and bark extensively while females often were found under oak leaves. For over 50% of the observations snakes were near blueberry or pitch pine, although males were near blueberry twice as frequently as females. There were no sexual differences in activity patterns, but individuals differed in habitat use and activity. Some snakes remained in only one habitat, whereas others moved between habitats. Descriptions of habitat use are numerous for reptiles (e.g., Heatwole, 1977), but individual differences have seldom been quantified. Most studies of habitat use by snakes provide quantitative descriptions of where the snakes generally occur (Wright and Wright, 1957; Reinert and Kodrich, 1982; Weatherhead and Charland, 1985; Burger and Zappalorti, 1986) but do not compare where the sexes are. Reinert (1984a, b) performed a detailed analysis of habitat use by rattlesnakes (Crotalus horridus) and copperheads (Agkistrodon contortrix mokeson), but the topics of sexual or individual differences in habitat use by snakes remain essentially unstudied. In this paper we examine sexual and individual differences in habitat use and activities of 10 radiotracked pine snakes (Pituophis m. melanoleucus) in the New Jersey Pine Barrens. Our goal was to obtain detailed data on how individuals use habitat, whether males and females inhabit the same habitats, and if there is potential for resource competition between the sexes. Further, we examine the relative importance of specific habitat features for individuals and show that a feature may be important to one individual but not to others. Pine snakes occur in a variety of habitats in North America, but are nowhere common (Wright and Wright, 1957). In the New Jersey Pine Barrens, pine snakes occur in pitch pine (Pinus rigida) and scrub oak (Quercus marylandica) woodlands, but avoid white cedar (Thuja occidentalis) swamps and lowland pitch pine areas (Zappalorti and Burger, 1986). Thus, snakes generally choose moist, but not wet ground cover and some vegetation that provides shade (Burger and Zappalorti, 1988). METHODS AND STUDY AREA We studied pine snakes near Toms River, Ocean County, New Jersey in 1981 and 1982. The study area contained residential commuce competition between th sexes. Further, a ine the relative importance of spe ific 68 This content downloaded from 207.46.13.13 on Fri, 26 Aug 2016 06:17:05 UTC All use subject to http://about.jstor.org/terms HABITAT USE IN SNAKES nities and forested sections of pitch pine, scrub oak and white cedar. Ten snakes (5 males, 5 females) were located soon after emergence from hibernation in early spring (March, April) in 1981 (4 snakes) or while nesting (June-early July) in 1982 (6 snakes). Snakes were sexed by everting hemipenes (Gregory, 1983), measured (SVL), and weighed. Radio transmitters (AMV Instrument Company SM 1; 3 g, 2.5 cm long) were inserted surgically in the body cavity, following procedures in Reinert and Cundall (1982) and Burger and Zappalorti (1988). Prior to implantation in the study animals, transmitters were surgically implanted in other pine snakes that were then observed for two weeks to ascertain normal behavior. The study area was searched 3-4 times a week from June through December in 1981 and 1982. Transmitter batteries did not last through the winter, and male 2 disappeared three weeks after release. When a snake was located, the following data were collected: forest type (Forman, 1979), elevation, soil type (U.S.D.A., 1980), disturbance type, nearest vegetation (over 5 cm high), nearest ground cover, distance to water, cloacal temperature, air temperature (sun and shade) and snake activity. Forest type, nearest vegetation and soil type were not necessarily correlated since the same vegetation may grow on different soils. Vegetation type and ground cover type were noted for each snake. In this area of the pine barrens, pine trees are sometimes sparse, and other vegetation is scattered between the trees. We distinguished four forest types: (1) oakpine (over 50% oak), (2) pine-oak (50-80% pine), (3) pine lowland (> 80% pine), and (4) cedar bog (no pine). We recorded soil types (Atsion, Berryland, Lakehurst, Lakewood; U.S.D.A., 1980) because we suspected females might be found on a particular soil suitable for digging nests. A site was considered disturbed if it showed evidence of past human activity (clearing for agriculture, hunting lodges, railroads, or logging roads), undisturbed if it did not. Snake activity was recorded as: (1) inactive (in the shade or hidden), (2) basking in the sun, (3) basking in partial shade, and (4) moving. We defined basking as a snake thermoregulating on a clear day with 50% or more of its body exposed to direct sun (basking in sun), or with less than 50% exposed to direct sun (basking in shade; after Parker and Brown, 1980). Wilcoxon X2 tests and Contingency Table X2 tests were used to determine differences in the distribution of variables among groups, and stepwise multiple regression procedures with maximum R selection criterion were used to identify the environmental variables that contributed to the variation in the dependent variables (Barr et al., 1976). The procedure determines the model giving the highest R2 with the fewest variables, and gives the R2 and F values for the contribution of each variable to the dependent measures. The model's procedure eliminates the effect of one variable (e.g., date), before examining the effect of another variable. A variable that varies colinearly with another is not entered in the model, and these are not presented.

We examined eight summer dens (used only in summer) and seven hibernacula (occupied both in winter and summer) of the snake Pituophis melanoleucus in the New Jersey Pine Barrens, comparing above ground characteristics of hibernacula and summer dens with characteristics at nearby random points. Temperatures at the soil surface and at 10 cm depth were significantly warmer, and there was less leaf cover around the random points compared to the entrances of the hibernacula and summer dens. Hiber- nacula had significantly more vegetation cover within 5 m, more leaf cover over the burrow entrance, and were closer to trees than were summer dens. Most hibernacula and summer dens were beside old fallen logs (73%), the entrance tunnels following decaying roots into the soil. Excavation of the hibernacula and summer dens indicated that most hibernacula appeared to be dug by the snakes and had an average of eight side chambers and 642 cm of tunnels, compared to less than one side chamber and 122 cm of tunnels for summer dens. Except for hatchlings, most snakes in hibernacula were located in individual chambers off the main tunnel; all snakes were at depths of 50-111 cm (X = 79 cm). Pine snakes may select optimum

Pine Barrens treefrog (Hyla andersonii) in Okaloosa County, Florida, 21 July 1982, I heard a distinctive but unfamiliar frog call. Investigation disclosed a small Rana sitting in shallow water along a clear stream. Through the remainder of the evening, similar calls were heard at five additional sites, and five more specimens were collected. It soon became apparent that these frogs represented an undescribed species. A total of 34 adults, three recently transformed juveniles and numerous tadpoles are now available. Specimens have been deposited in the Florida State Museum (UF) and the United States National Museum (USNM).

Range extensions for the eastern hognose snake, Heterodon platirhinos, are compared to known and new data from eastern New England: Rhode Island, Massachusetts east of the Worcester County line, and New Hampshire. The species is unknown from coastal islands; reasons are suggested. The disjunct New Hampshire population occupies inland sandy habitats dominated by white pine (Pinus strobus). Albino and melanistic specimens occur. Eastern New England, including New Hampshire, Rhode Island, and Massachusetts east of the Worcester County line, is regarded as part of the peripheral range of Heterodon platirhinos, where the species is "rare or only locally common in semi-isolated populations" (Platt, 1969). The only previous evidence we can find for occurrence of this species in New Hampshire, for example, is the photograph of an albino and the letter published by Newton (1940). The species is not, however, listed by the state of Massachusetts for special consideration (Blodget and Cardoza, 1983). New Hampshire has no state listings at present. One of us (JDL) has extensive experience with this species in the southern region of eastern New England, and the other (MCM) in the northern portion. Our field experience overlapped for many years in the centrally located suburban Boston area. Because Edgren (1961) examined no New Hampshire specimens, and saw only 25 from Massachusetts, we believe a range and population analysis is warranted at this time. Aspects of geographic distribution are shown in Fig. 1. Little (1971) plots eight major environmental variables of climate and physiographic features. None corresponds closely with the limits of hognose snake distribution. Only the ranges of two tree species, swamp white oak, Quercus bicolor, and Atlantic white cedar, Chamaecyparis thyoides, show some correspondence (Little, 1971). Both of these trees are characteristic of wet areas in typical southern New England pine barrens, preferred by the hognose snake in much of its range, where adjacent drier areas are dominated by pitch pine, Pinus rigida. Our records from southern New Hampshire (Hillsborough and Merrimack counties) show good correspondence to inland soils derived from sandy outwash material (Goldthwait et al., 1951), but not with the sandy coastal plain soils utilized by these snakes from Long Island southward (Conant, 1975). We have previously (Lazell and Michener, 1976) suggested that three factors control hognose snake distribution in the sandy soils of this region: 1. Heterodon platirhinos is a southern form, whose range has only recently moved north; therefore, the species arrived in New England after sea levels had risen significantly since the Wurm (ca. 10,000 yr BP), and the species has not colonized across sea water. 2. These snakes must have an abundance of toads (Bufo spp.) as prey to sustain adult populations. 3. They must have a supply of small amphibians, either salamanders, Plethodon cinereus, or peepers, Hyla crucifer, to sustain hatchlings and young until they are big enough to eat toads. Small toads alone cannot supply the young snakes' diet, because these amphibians do not metamorphose into the terrestrial form until well into the summer of each year. Hognose snakes are thus able to colonize peninsular habitats where soils are loose enough to permit burrowing and the habitats can supply needs two and three above. Hognose snakes seem to be absent from all of the islands in Rhode Island (Table 1). Similarly, hognose snakes are absent from all of the Massachusetts coastal islands including Monomoy, which has been connected to the mainland and which does support toads. Monomoy lacks both woodland salamanders and peepers (Lazell and Michener, 1976). Two peninsular barrier systems support good populations of hognose snakes: Sandy Neck and the Provincelands, in Barnstable County, Massachusetts. Details of their physiography are given in Lazell (1979). A mark recapture survey has been begun on Sandy Neck; preliminary results indicate approximately one hognose snake per three hectares of land, excluding salt marsh. Hognose snakes are abundant north of Provincetown in Provincelands, where at old This content downloaded from 157.55.39.251 on Fri, 13 May 2016 07:23:39 UTC All use subject to http://about.jstor.org/terms M. C. MICHENER AND J. D. LAZELL FIG. 1. Distribution of the hognose snake in eastern New England. Black circles are museum specimens, open circles photographs; the number of individuals in each case is indicated if more than one. Sight records are indicated by X. Urban areas are stippled. dump sites one may find more than one per hectare. Both of these peninsulas supply all three range requirements noted above. We are convinced that Heterodon platirhinos, albeit more or less common, is fairly continuously distributed south of Conant's (1975) line, given minor adjustments (Fig. 1). Because this snake does well in edificarian habitats, and is most common in the pine barrens ecosystem TABLE 1. The major islands of Rhode Island. None is known to support hognose snakes. Habitat characteristics are more fully described in text. Habitat prerequisites

Radiotelemetry was used to monitor the movements and habitat use of timber rattlesnakes (Crotalus horridus) in the Pine Barrens of southern New Jersey. Reproductive condition strongly influenced both aspects of behavior. Males generally exhibited the largest activity ranges, and the sizes of their ranges were positively correlated with the number of days the snakes were monitored. This was not true for gravid or non-gravid females. Time series analyses indicated that movement patterns of males and non-gravid females consisted of constantly shifting, non-overlapping activity areas. In most cases these snakes moved in a looping pattern during the active season that returned them to the same hibernation site from which they departed. Gravid females exhibited more static, overlapping activity areas and shorter dispersal distances from hibernacula. Males and non-gravid females utilized forested habitat with greater than 50% canopy closure, thick surface vegetation (approx. 75%), and few fallen logs. This habitat occurred with high frequency throughout the study area. Gravid snakes utilized less densely forested sites with approx. 25% canopy closure, an equal mixture of vegetation and leaf litter covering the surface, frequent fallen logs, and warmer climatic conditions. This habitat occurred in low frequency on the study area and was largely restricted to the edge of sand roads.

ON June 9, 1929, I found a small frog hopping about in grassy intermediate pine barrens on Trail Ridge, about 4 miles west of Folkston, Charlton County, Georgia. Through some rather unaccountable lapsus I took it at the time for a Hyla femoralis, and proceeded to pose it on a near-by pine for a photograph. I recall that it did not cling very well to the vertical trunk, but fell off several times before the photograph (fig. 1) was secured. I must have been at least dimly aware of something unusual about the specimen, for I took occasion to make note of yellow spots on its femora and prominent black blotches on its sides.